Modern Textbook Of Zoology: Vertebrates 8171338917, 9788171338917, 9788171338, 9789350440414, 9350440415

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MODERN TEXT BOOK OF ZOOLOGY

v

T

[ ANIMAL DIVERSITY - II ]

R.L.KOTPAL FORMERLY, PROFESSOR AND HEAD DEPARTMENT OF ZOOLOGY

MEERUT COLLEGE MEERUT

MODERN TEXT BOOK OF ZOOLOGY

[ANIMAL DIVERSITY - II]

NEW DELHI OFFICE: 9, R AN I ]H ANS I ROAD ( MO TI A KH AN) NEW DELH I 11 0055

MODERN TEXT BOOK OF ZOOLOGY

VERTEBRATES ISBN 81-7133-891-7

ISBN No. : 978-81-7133-891-7 © RESERVED

AI/ rights reserved. No part of this book (allY editioll/reprint) may be produced, stored in a retrieval system or trallsmitted in anv jorm what so ever or by an)' means electronica//), or mechanical/\, or bv photocopying, recording or otherwise without the prior \I'ritten permission of'the Publisher In!i'ingement o/copyright is a criminal o/fence, TITLE CODE NO.

Z-3

EDITION • 2009

PUBLISHED

BY

RAKESH

KUMAR

- 21nO

RASTOGI

PUBLICATIONS. 'GANGOTR I' SHIVAJI

ROAD.

PHONES. 0121 - 2510688. 2515142. 2516080 email. [email protected]

FOR

RASTOGI

MEERUT-250 002 FAX.

0121 - 25215~5

Websi le. ,I\\\\ .r" ' (02 Ipublicallnll , .cllm

PRINTED AT CAPTIAL OFFSET PRESS

NEW

DELHI.

INDIA

2 (DIFFERENCES)

CHORDATES

NON-CHORDATES. ORIGIN MAJOR GENERAL,

.HAKi'"

CHORDATA.

CL.AS~>lFlCI'\TIO'N OF CHORDATA WITH CHARACTERS.

NON-CHORDATES. OF

LHUr\.'Un..,rI

OVER OTHER PHYLA

Contents

1 SYSTEMATIC POSITION OF (HEMICHORDATA).

OTHER HEMICHORDATES . DOUCHOGLOSSUS). 4. CEPHALOD1SCUS.

SYSTEMATIC POSITION OF HERDMANIA (UROCHORDATA).

, GENERAL CHARACTERS.

OTHER UROCHORDATES

CLASSIFICATION. PYROSOMA.

DOUOLUM.

PTYCHODERA ATUBARIA.

Contents

GENERAL CHARACTERS. PRIMITIVE, DEGENERATE AND

AFFINITIES

SYSTEMATIC POSITION.

SPECIALIZED CHARACTERS OF BRANCHIOSTOMA (CEPHALOCHORDATA).

9 WHAT ARE PROTOCHORDATES SIGNIFICANCE OF PROTOCHORDATES.

COMPARISON OF THREE PROTOCHORDATE SlJBPHYLA GENERAL COMPARISON.

HABITS AND

3. ALIMENTARY CANALS AND ASSOCIATED GLANDS. 4. PHARYNX. 5. FOOD AND FEEDING. SYSTEM. 7. NERVOUS SYSTEM).

WHAT

VERTEBRATES

PHYLOGENY

OR

EVOLUTIONARY

CHORDATES VERSUS VERTEBRATES.

VERTEBRATES.

GENERAL CHARACTERS OF SUBPHYLUM VERTEBRATA.

ORIGIN AND ANCESTRY OF VERTEBRATES

OF

HISTORY

DIVERSITY OF VERTEBRATES.

1.

11

OCCURRENCE.

BIOLOGICAL SIGNIFICANCE.

IMPORTANT FEATURES.

INTERRELATIONSHIPS AND AFFINITIES.

CLASSIFICATION.

11

1

x

1

Contents

1 DIFFERENCES BETWEEN

GENERAL CHARACTERS.

31

HAGFISHES.

AFFINITIES OF CYCLOSTOMATA. PHYLOGENETIC STATUS OF CYCLOSTOMATA.

CYCLOSTOMES BDElLOSTOMA}.

1

3 CLASSIFICATION. BIOLOGICAL SIG,Nn:'1C}~NC:E

THE FIRST JAWED OCCURRENCE. IMPORTANT FEATURES.

1 ECONOMIC IMPORTANCE.

1 HEPTRANCHlAS,

GENERAL CHARACTERS. CLASSIFICATION. OTHER CHONDRICHTHYES . DOGFISHES (SCYUQRHINUS, SQUAWS, SHARKS (RHINCODON. CEfORHINUS, CARCHARODON. CARCHARINUS,

HEXANCHUS,

MYLIOBATIS. DASYATIS. TORPEDO).

AND

GUITARFISH (RH1NOBATUS). CHIMAERA).

CHIMAERAS

CHLAMYDOSELACHUS.

1 ECONOMIC IMPORTANCE.

Contents

5. GENERAL

16 UI':O.''''''''''. CHARACTERS

OF SUPERCLASS ACCESSORY

BETWEEN

CHONDRICHTHYES

1:,

PHOTORECEPTORS

In

1

xx J

Contents

81 I. HABITAT, DISTRIBUTION

HABITS.

EXTERNAL FEATURES. INTEGUMENT AND EXOSKELETON.

4. SKULLS. PECTORAL

6. PELVIC

DIGESTIVE SYSTEM. RESPIRATORY SYSTEM. HEARTS. BRAIN.

12. URINOGENITAL SYSTEM.

",,,'d,.fe,,"-.

LIMB

1

is

CharacteJrs Com,moln to Chc)rd:ne to a more potent androgen called dihydrotestosterone (DHT). FSH and LH act together on sustentacular cells (Sertoli cells) which produce a protein, the androgen binding protein (ABP) which binds to testosterone which keeps the concentration of testosterone high near the tubules. Testosterone stimulates the final steps of spermatogenesis. Figure explains the various events more elaborately.

HEAD

centriole

MIDDLE PIECE

cell membrane TAIL

8

Oogenesis

(Z-3)

mature sperm interstitial cells cell sertoli cells

Fig. 58. Cross section of seminiferous tubule of man showing spennatogenesis.

Hormonal control of spermatogenesis

It is the process through which the haploid (h) secondary oocytes are formed inl the ovaries. It includes several phases including meiosis.

.

A Fig. 59. Structl!I'C of mammalian sperm under electron microscope. A-Complete spenn, B-A part magnified.

Oryctolagus

The Rabbit

r--f I

I I

LRH (LH releasing hormone)

"-

/

I inhibits LRH secretion LH(ICSH)

stimulates spermatogensis?

n~d~~r spermatogenesis

bone growth

o

first polar body

secondary oocyte

sperm cell

I \

-+ beard lower voice

Fig. 60. Reproductive hormone relationship in the male, dotted line shows inhibition.

Reduction division (Meiosis I) During early development of the foetus the primordial germ cells migrate from the endoderm of yolk sac to lie in the ovaries and differentiate into deploid (2n) oogonia. The oogonia divide mitotically to give rise to a large number of germ cells. Many of these germ cells undergo degeneration and a few develop into primary oocytes which enter into prophase of reduction division (meiosis I) during fetal development but do not complete it till puberty. Each primary oocyte is surrounded by a single layer of follicular cells, and the entire structure is called the primordial follicle. The primary follicles are surrounded by a single layer of cuboidal cells. Later on more layers are added to it (6-7) called the granulosa cells. A glycoprotein layer zona pellucida develops between the primary oocyte and the granulosa cells. The

zygote

the secondary oocyte begins equatorial division (meiosis II) but stops in metaphase

0

a secondary oocyte (and firsl polar body) is ovulated

secondary ...--'-----. oocyte

~"0 + G

after puberty, primary oocytes complete meiosis I, which produces a secondary oocyte and a first polar body that mayor may not divide again

0

fertilization

/!~

maturation ot reproductIVe structures

I ~®

\ I I I I

TESTOSTERONE

primary oocyte

~ion0

stimulates interstitial cells

+--

G

f

I~---T--~~==~--~--~ TESTES

I I

t

I I

during fetal development reduction division (meiosis I) begins but stops in prophase

oogonium

I

I FSH

G

--1 I

hypothalamus

FRH (FSH-releasing hormone)

inhibiting substance not yet identified (inhibin? estro.gen?)

[ 483

second polar body

after fertilization, equatorial division resumes. The oocyte splits into an ovum and a second polar body.

The nuclei of the sperm cell and the ovum unite, forming a diploid (2n) zygote.

Fig. 61. Summary of events associated with fertilization and implantation.

innermost layer of the cells becomes firmly attached to the zona pellucida and is called the corona radiata. The outermost granulosa cells form theca folliculi which consists of the theca interna and theca externa. The granulosa cells secrete a fluid which accumulates in the cavity of the follicle called antrum. The follicle is now called the secondary follicle. After puberty, under the influence of the gonadotropin hormones secreted by the anterior pituitary gland meiosis starts in some of the follicles. The diploid primary oocytes complete reduction division and two haploid cells of unequal size are formed both with half number of chromosomes. The smaller cell so produced is called the first polar body. The polar body contains mostly the discarded nuclear material and very little cytoplasm. The larger cell known as the secondary oocyte proceeds to the metaphase of the equatorial division (meiosis II) and then stops at this stage. The follicle containing such oocytes are

Oryctolagus

484 J

The Rabbit

embryo implants in uterus

menstruation does not occur "---v---J

endometrium of uterus Fig. 62. intenuption of menstrual cycle after pregnancy in human.

cells tissue }

ovarian ligament

f~

0-~·-----sE'condarv oocyte

0=

~

.2

J:u:'!!~--' --'''''',.}l\>.f~ elongated snout

--~

small pinna -----.llin

Fig. 10. Flying mamlllais or bats. A-An insect-eating bat (Microciuroptera). B-Head of a vampire bat, Desmodus. C-A truit-eating bat (Mcgachiroptera).

radial sesmoid or falciform bone. Moles are of some commercial value because of their fur and because of the insects they destroy. 3. Erinaceus (Hedgehog). Hedgehogs range throughout Europe, Africa and Asia. They have a small globular body, pointed snout, stumpy tail and short legs. Skin is covered with short barb less spines, intermingled with hair, except on the belly. When disturbed, they roll up into a ball for protection, so that the spines project outward like pins in a pincushion. They are omnivorous creatures, feeding on insects, snails, young birds and snakes. It lives under bushes or In subterranean channels. Common Indian name is 'Jhau Chuha'. Common Indian genera are Erinaceus, Hemiechinus (Rajasthan) and Paraechinus (South India). [IV] Flying mammals

Order Chiroptera includes bats which are the only true flying mammals that compete very weII with the birds. Their forelimbs are modified into wings or patagia. These are the membranous leathery extensions of skin along the two sides of body, limbs and tail. The wings are supported by greatly elongated forearms and four fingers (2nd to 5th). The hindlegs are small and have clawed digits by which they hang upside down to a tree branch or perch, while sleeping during day. All are nocturnal

with remarkable power to avoid obstacles while in flight at night. They have a highly developed sonar or echoapparatus, a kind of radar. Their vocal cords produce ultrasonic sounds, inaudible to human ears. When striking objects, their reflected echos are caught by the highly sensitive ears, and utilized to avoid objects during flight. Bats may be frugivorous, insectivorous or sanguinivorous (Table I, Fig. 10). 1. Pteropus or Cynopterus (Frugivorous bats). These are the fruit-eating bats found in the Old World tropics and sub-tropics, including India. They belong so the suborder Megachicroptera. They are called the flying faxes because of fox-like head, long snout and large eyes. Some of them are the largest of bats with a body 30 cm in length and a wingspread of 1.5 metres. Body is covered with brown fur. Molars are marked with a longitudinal groove and are not tubercular. They have short rudimentary tails and a claw on the second as well as first finger which is free from patagium. 2. Rhinolophus (Insectivorous bats). Insectivorous bats belong to the suborder Microchiroptera. They are small to medium in size, with small eyes, comparatively short snout and without claw on second finger. Most have nose leaves and special flaps in front of their large ears, used for navigation or echo-location.

502 j

Class Mammalia

Common examples are horseshoe bat (Rhinolophus), mousetailed bat (Rhinopoma), etc. 3. Desmodus (Vampire bat). Tree vampire bats occur in tropics and are famous because of their sanguinivorous habit. They often feed on fresh blood of sleeping domestic animals, even human beings. Their incisors are razor-sharp with which they slit the skin of the prey, where hairs and feathers are scanty, and the oozing blood is lapped up. The vampires are very sly so that the sleeping victim often remains unaware. They transmit paralytic rabies in their victims. Associated with the blood-sucking habit is the very small lumen of their oesophagus through which solid food cannot pass. The first digit of the forelimb bears claw. Tail is present. Pinna are large with a lobe called tragus. Molars bear transverse groove. Vampires do not have a nose-leaf. Instead, they have a naked pad wearing U-shaped grooves. Common genera are Desmodus and Megaderma. [V] Toothless mammals Toothless mammals (Order Edentata) are highly specialized and rather primitive New World mammals. They are either toothless or else have

degenerated teeth, without enamel. Ants form a large part of their diet. They include sloths, anteaters and armadillos (Fig. 11). 1. ldadypus (Sloths). Sloths of tropical America are distinctly arboreal and slow-moving creatures. They have short rounded heads, inconspicuous ears, fflrward-Iooking eyes and no tails. ThlJ hands and feet bear 2 or 3 long and curved claws by which the animal hangs upside down from tree branches, eating, sleeping and moving in that position. There are no incisors, but molars are present. In the neck there are 9 cervical vertebrae and excessive number of trunk vertebrae. Algae often grow on their straw-like hairs providing a green protective colouration among the leaves. They feed mainly on fruits and leaves. The 3-toed sloth is Bradypus tridactylus, while the 2-toed slots is Choloepus didactyl us. 2. Myrmecophaga (Great anteater). Anteaters are characterized by long tapering snouts, tubular mouths, without teeth and long sticky protrusible tongues for capturing ants and termites which are swallowed whole. Their forefeet have long, sharp claws for digging open the anthills. Hind limbs have 4 digits with normal claws. While walking r

transverse bands of bony scutes

non-prehensile tail Three-toed sloth Bradypus tf/dactylus

Cape anteater or aardvark Orycteropus afer

Fig II. Toothless mammals and anteaters.

Pangolin or Scaly anteater ManiS tn'CUSPIS

[ 503

Class Mammalia digits of manus are bent and their dorsal surface is placed on the ground. There are three living anteaters-(i) giant (Myrmecophaga tridactyla), about 2 metres long, with a bushy tail, (ii) lesser (Tamandua tetradactyla) about 60 cm long, with a nonbushy prehensile tail, and (iii) pigmy anteater. 3. Dasypus (Armadillo). The nine-banded armadillo (Dasypus novemcinctus) found in South and Central America, is about 75 cm long. It has a well-developed dermal skeleton of transverse bony scutes. These are covered with horny plates or scales with hairs embedded and connected by flexible skin. In adults incisors and canines are absent but there are numerous continuously growing molars with no enamel. When threatened, it rolls up into a ball for protection. Strong claws on front feet are used for burrowing. The animal shows an interesting case of polyembryony in which a single fertilized egg produces 4 to 8 young ones of the same sex.

[VI] Other anteaters The Old World counterparts of the New World edentates are the scaly anteaters or the pangolins (order Pholidota) and the cape anteater or aardvaek (Order Tubulidentata). Their superfi(;ial resemblance with edentates is probably due to similar mode of life (convergent evolution). 1. Manis (Pangolin). Scaly anteaters or pangolins have their home in tropical Asia and Africa and belong to the single genus Manis. They live on ground, in burrows or in trees and range from 30 cm to 1.5 metres in length. Their long tapering body is covered with large overlapping epidermal scales, except on snout, sides of face and undersurface. They have a long, sticky, extensile tongue with muscular roots, used for feeding on termites or white ants. Stomach is reduced and linked with as thick keratinous layer for grinding insects. The 5 strong front claws are used to tear open termite's nests and for burrowing. When attacked, they roll themselves into a ball for defence, like armadillos. Common Indian specIes are Mallis crassicuudata (South

India), M. pentadactyla (India to Sri Lanka) and M. aurita (Nepal to China). 2. Orycteropus (Cape anteater). Order Tubulidentata is represented by a single genus (Orycteropus) with 3 or 4 species of aardvarks or cape anteaters widespread in African grass lands. Aardvark is the Dutch name meaning earth-pig. These peculiar animals have a 75 cm long pig-like body, long donkey-like ears, an elongated snout with a tubular mouth and wide nostrils, and a long heavy tail. Tongue is long, sticky and extensile for collecting ants and termites which form their principal diet. With strong claws they are expert diggers.

[VII] Gnawing mammals Gnawing mammals are mostly small in size and characterized by the absence of canine teeth, and great development of incisors which continue to grow throughout life and used in gnawing. They belong to two orders : Rodentia (rats, mice, squirrels, chipmunks, gophers, porcupines, beavers, woodchucks, guinea pigs, hamsters, chinchillas, etc.) and Lagomorpha (pikas, rabbits, hares). True rodents have only I pair of chisel-shaped incisors in each jaw, whereas lagomorphs have a reduced second pair besides the functional pair of upper incisors. The two orders seem to resemble due to parallel evolution. Gnawing has been a very successful mode of life since rodents are found all over and exceed in number of species and individuals than all other mammals combined (Fig. 12). 1. Rattus (Rat). Rats are the most harmful and well-known rodent pests, living in holes and burrows in the houses and in cultivated fields all over the world. They have naked, scaly tails and long snouts. They are very cunning, gregarious. nocturnal and prolific breeders. They are host to ratHeas which carry the bacillus Pasteurella pestis causing bubonic plague in man. Cr-i- alisphenoid

posltrontal

sphenotic

preopercular ....,,~~+-

hyomandlbular

prootic

opercular supratemporal exoccipital

parietal

transverse process of 1st vertebra

channel for dorsal aorta

B

Fig. I. Labeo. Skull. A-Dorsal view. B-Ventral view.

posteriorly by sphenotic. Besides, each eye is encircled by a chain of 5 orbital bonessupraorbital, postfrontal, postorbital, infraorbital and preorbital. (vi) the nasal or ethmoidal region comprises 9 bones of nasal capsules and snout. Paire4 bones one on either side are nasals, ectoethmoids and lacrymals. Median single bones are mesethmoid (dorsal), rostral (anterior) and vomer (ventral). 2. Visceral skeleton. It includes 7 paired arches corresponding to those of the dogfish. (i) The first or mandibular arch comprises two parts. The upper part (palatoquadrate bar) attaches to cranium forming the upper jaw, each half of which includes two membrane bones (premaxilla and maxilla) and three replacing bones (palatine, metapterygoid and quadrate). The lower part (Meckel's cartilage) forms the lower jaw, each half of which includes three bones-dentary, angular and articular. Articular hinges on quadrate which attaches to cranium. All the jaw bones lack teeth. (ii) The second or hyoid arch includes the upper suspnsorium including two large bones (hyomandibular and symplectic), and a lower hyoid

..,.....c+-- opercular

Interopercular

Fig. 2. Labeo. Opercular bones.

cornu which supports the tongue and floor of bu(;cal cavity. Investing bones or bony plates supporting the operculum on either side are : opercular; preopercular; interopercular and subopercular (Fig. 2). (iii) Remaining 5 arches are called the brt..~.chial or gill arches. Each of the 3 to 6 arches bears gill lamellae on the outer border, and a double row of small spiny gill rakers on the inner pharyngeal border. The 7th arch is reduced on either side into a triangular i1lj''I"ior pharY1lgeal

552 ]

. Endoskeleton of Labeo rohita

Fig. 4. Labeo. Weberian ossicales and air bladder of a teleost fish in left dissection.

A

parapophysis

Fig. 3. Labeo. Vertebrae. A-Trunk vertebra in posterior view. S-Typical caudal vertebra in posterior view. C-First caudal vertebra in anterior view.

bearing three rows of strong masticating plates or teeth. Typically, a branchial arch consists on either side of four bones, supporting the pharyngeal wall. These are dorsal pharyngobrallchial, lateral epibranchial and ceratobranchial. and ventral hypobranchial which is connected to a common flat median basibranchial lying in the floor of pharynx.

Vertebral Column lThe vertebral column is welJ ossified and composed of 37-38 simple and comparatively similar vertebrae. 1. Trunk vertebra. A typical trunk vertebra consists of : (i) a cylindrical centrum deeply concave at both ends (amphicoelolls), (ii) a small dorsal neural arch around the spinal cord, (iii) a slender neural spine for muscular attachment, (iv) a pair of short transverse processes or parapophyses arising ventro-Iaterally from centrum, and (v) a pair each of small blunt facets on neural arch, called pre-and post- zygapophyses, by which adjacent vertebrae articulate. These articular facets do not occur in the vertebrae of elasmobranchs. Ribs are attached by ligaments to parapophyses of

trunk vertebrae and serve as a protective framework for the bodycavity and its contents (Fig. 3A). 2. Caudal vertebra. A typical caudal vertebra basically has the same parts as a trunk vertebra. But there are no ribs. Also, the centrum bears ventrally a haemal arch around the caudal artery and vein and projecting below into a long slender haemal spine (Figs. 3B & C). 3. Weberian ossicles. In cyprinoids (e.g. Labeo) and siluroids (order Ostariophysi), a chain of four small bones connects the airbladder and the internal ear on either side. These are named Weberian ossicles after the name of their discover (Weber, 1820), who regarded them to be ear ossicles. However, they are not homologous to the ear ossicles of other vertebrates as they are formed by the segments of few anterior trunk vertebrae. The four Weberian ossicles are claustrum, scaphium, intercalarium and tripus (Fig. 4). Neural arch of first trunk vertebra forms a small dorsal claustrum and another small ventral scaphium which is in direct contact with the internal ear. The centra of second and third trunk vertebrae are completely fused together. The transverse process of third vertebra becomes modified into the large triangular and anteroposteriorly extended tripus which is in contact with the anterior end of air bladder. Intercalarium is a small bone with a backwardly directed spinous process. It lies embedded in a ligament between scaphium and

[ 553

Endoskeleton of Labeo rohita lateral fused piece of neural arches 2 + 3

median piece of fused neural arches 2 + 3

neural spines of caudal vertebrae centrum

fused neural spines \.~_ .--rn - " 2+3 ~ __ ':.: ~.\.,:.\ \1",11, ,0'.' neural spine 1 ,~ :'

haemal spine

neural arch 1 '.

A radials intercalarium proximal piece of radial

mesial piece of radial

finrays (Iepidotrichia)

distal peice of radial

Fig. 7. Lllbeo. A-Skeleton of ventral fin and corresponding caudal vertebrae. B-Two radials and fin-rays of dorsal fin.

Fig. 5 Labeo. Weberian ossicles attached to vertebrae in left view_ intercalarium

scaphium

articulating process transformator

anterior ramus

body of trip us

Fig. 6. Lllbeo. Chain of Weberian ossicles separated.

tripus. The actual derivations and functions of Weberian ossicles remain controversial. They probably help in maintaining equilibrium by conveying changes of air pressures in the air bladder to the internal ears (Figs. 4-6).

Median Fin Skeleton The thick basal part of a median fin (dorsal or anal) is supported by a series of parallel, rod-like bones, the radials or somactidia, which remain

embedded in muscles. Typically, each radial or somactid is made of three segments : proximal, middle and distal. The proximal piece is large, dagger-shaped and attached by ligament to neural or haemal spine of corresponding vertebra. Middle piece is short and rod-like. Distal piece is very small and connected to the dermal, bony and branched slender finrays, called lepidotrichia, which support the membranous fin. At the free edges of fins are also present unbranched horny actinotrichia. In caudal fin, some radials fuse with vertebral neural spines to form epurals and with haemal spines to form hypurals. Caudal fin of Labeo has 2 epurals and 1 radial dorsally, and 9 hypurals ventrally. Lepidotrichia are arranged in 2 symmetrical halves (Fig. 7).

Pectoral Girdle and Fins 1. Pectoral girdle. It lies just behind the last branchial arch and consists of separate lateral halves. They do not meet mid-ventrally as in Scoliodon. Each half comprises an inner primary and an outer secondary part. Primary part is composed of 3 replacing bones : a ring-like scapula with a large scapular foramen, a large

554 J

Endoskeleton of Labeo rohita

supracleithrum

scapular foramen

pelvic fin

Fig. 8. Labeo. Left half of pectoral girdle with attached fin.

irregularly triangular and fenestrated coracoid. and a A.-shaped mesocoracoid. Scapula and coracoid provide the glenoid articulation to pectoral fin. Secondary part is composed of 4 investing bones : a small conical post-temporal. an elongated dagger-shaped supracleithrum. a large crescentic clavicle or cleithrum. and a stout curved postcleithrum. The post-temporal articulates with the pterotic process of skull (Fig. 8). 2. Per.toral fin. Each pectoral tin is supported by 19 finrays or lepidotrichia proximally joined with 4 radials or somactidia, 3 of which attach with glenoid articulation of scapula and coracoid.

Fig. 9. Labeo. Pelvic girdle and right pelvic fin in ventral view.

the midventral line. Each half is made of a single large replacing bone, the basipterygium or pelvic bone. Its anterior broad part with forked end is connected in front by a ligament to the rib of the 12th trunk vertebra, while the posterior narrow rod-like part tapers behind into a small cartilage and also unites with the fellow of the other side in the mid-ventral line (Fig. 9). 2. Pelvic fin. Each pelvic fin is supported by 9 lepidotrichia, (fin rays), 3 somactidia (radials) and a supernumerary fin ray.

Pelvic Girdle and Fins 1. Pelvic girdle. It lies in the ventral abdominal wall anterior to anal fin. Its two halves meet in

IMPORTANT QUESTIONS

»

Shon Answer Type Questions I 2. 3

Draw labelled diagrams of dorsal and ventral views of skull of Labeo. Descnbe the structure of typical trunk or caudal vertebra of Labeo. How does it differ from other vertebrae ? Give the structure of pectoral or pd\'lc girdle of Labeo.

Endoskeleton of Labeo rohita

» I.

[ 555

MuUiple Choice Questions

The advancement of axial skeleton of Labell over that of

7.

SClIlilltilm is the presence of :

2.

3.

4.

5.

6.

(a) Ribs (b) Skull (c) Vertebral column (d) Girdles The visceral skeleton in Labell posses~es how many pairs of arches: (a) 6 (b) 7 (c) 8 (d) 9 Median occipital spine is present in : (a) Exoccipital (b) Basioccipital (c) Supraoccipital (d) ProotiC bone The roof of cranium in Labell is fonned by : (a) Parasphenoid (b) Alisphenoid (c) Frontals (d) Parietals and frontais On the dorsal side of orbits in Labeo lie the : (a) Frontal (b) Alisphenoid (c) Orbitosphenoid (d) Ectoethmoid The unpaired bone of nasal region In Labell is : (a) Nasals (b) Mesetlunoid (c) Ectoethmoids (d) Lacrymals

8.

9.

The membrane bone of upper jaw in Labell : (a) Palatine (b) Metapterygoid (c) Premaxilla (d) Quadrate The visceral gill arches called as branchial arches are : (a) 2nd to 6th (b) 3rd to 7th (c) 4th to 8th (d) 3rd to 6th The number of vertebrae in the vertebral column of Labell :

(b) 38-39 (a) 37-38 (c) 39-40 (d) 40-41 10. Trunk vertebra in Labell is : (a) Procoelus (b) Amphicoelus (c) Acoelus (d) Opisthocoelus II. Bones connecting air bladder and internal ear are collectively known as : (b) Claustrum (a) Tripus (c) Weberian ossicles (d) Ear ossicJes 12. Each pelvic fm is supported by how many fin rays: (a) 7 (b) 8 (c) 5 (d) 9

ANSWERS 1. (a) 2. (b) 3. (el 4. (d) 5. (a) 6. (b) 7. (c) 8. (d) 9. (a) 10. (b) 11. (c) 12. (d)

Endoskeleton of Frog The living Amphibia do not possess an exoskeleton. The endoskeleton of frog is made partly of cartilage and partly of bOlle. In the early stages of development (tadpole), the skeleton is solely cartilaginous but, in the adult frog, it IS greatly replaced by bones called the cartilage or replacing bones. Bones become also developed in other parts of skin or dermis of certains regions where there was no pre-existing cartilage. Such bones are termed membrane bOlles. A part of primary cartilaginous skeleton is I mpregnated by calcium salts and becomes hard. This is known as the calcified cartilage. As usual, the skeleton is conveniently divided into axial and appendicular ~keleton. Axial skeleton includes skull, vertebral column and sternum which lie along the medIan longitudinal axis of body. Appendicular skeleton comprises the skeleton of the limbs and the girdles supporting them (Fig. 1).

Characteristic Features of Skull Skull of frog is characterized by the following important characters : (1) It is triangular in shape, broad and dorso-ventrally flattened. (2) A considerable part of chondrocranium of tadpole persists in the adult. A large part of cranium and sense capsules consists of cartilage. (3) Cranium is comparatively small and narrow due to small size of brain which it encloses. (4) Occipital region is greatly reduced. (5) Skull is dicondylic, i.e., it articulates with the atlas vertebra by two occipital condyles, one on each exoccipital. (6) Basi-, ali-, orbito-, and pre-sphenoids and supra- and basi-occipitals are absent. (7) Skull is platybasic because an interorbital septum is absent so that cranium reaches forwards, uninterrupted in the orbital region.

Endoskeleton of Frog

[ 557 maxilla

premaxillae

nasal SKULL

suprascapula

PECTORAL GIRDLE

FORELIMB BONES

VERTEBRAL COLUMN

PELVIC GIRDLE

Fig. I. Frog. Complete skeleton in dorsal view.

(8) Tabular sphenethmoid forms the posterior wall of olfactory chambers. (9) Vomers bear vomerine teeth. (10) Floor of cranium is formed by a large, dagger-like parasphenoid bone, whereas the roof is formed by Jronto-parietals. (11) Jaw suspensorium is autostylic, i.e., the lower jaw is attached to skull through a rod-like cartilage, the quadrate. (12) Tympanic ring-like. Tympanic bulla absent.

Parts of Skull The skull includes 3 parts : Cranium, sense capsules and visceral skeleton (Figs. 2-7). [I] Cranium

It encloses the brain, hence the name brain box. It is mostly cartilaginous (Fig. 2). 1. Occipital segment. It is the posterior-most part of cranium or brain-box. When detached from

Endoskeleton of Frog

558 J premaxillary teeth -:;::;:~it'ti'~:o..... maxillary teeth ---..~

septomaxillary

'-"'--=- cartilaginous nasal capsule -~~~ diamond-shaped area of sphenethmoid

vomerine teeth

or olfactory chamber

\

~

l~~:iili\-J,~ t:::::;=·~,

palatine

~~=::~~~~ ~~~~~~Iforamen for

maxilla

2:

pterygoid

V a nerve

w--~~~..

foramina for II nerve

~:;;~Pi~~:-i\\- VI nerve V& VII

toramen magnum

exoccipital

OCCipital condyle

fenestra ovalis

B Fig. 2. Frog. Skull. A-Dorsal view. B-Ventral view. frontoparietals

foramen magnum

prootic

auditory capsule

occipital condyles

fenestra ovaUs

(Dorsal view)

OCCIPITAL SEGMENT

(Dorsal view)

A

8

(Ventral view)

(Dorsal view)

PARASPHENOID

D

(Veneral view)

C

SPHENETHMOID

(Dorsal view)

(Ventral view)

SEPTOMAXILLARY

E

(Ventral view)

FRONTOPARIETALS

(Dorsal view) (Ventral view) NASAL

F

VOMER

G

Fig. 3. Frog. Loose skull bones of cranium and sense capsules.

skull, it appears somewhat like a vertebra. It contains a large hole, the foramen magnum, through which spinal cord enters cranium. Two irregular cartilaginous bones, called exoccipitals, lie one on either side of the foramen and almost completely surround it. Exoccipitals bear, on their posterior surfaces, two large oval convexities, the

occipital condyles, which articulate with the anterior two concavities of first vertebra or atlas. A cartilaginous auditory capsule is fused fmnly on the outer side of each exoccipital, forming a wing-like projection. A minute opening, fenestra ovalis, leads into the auditory capsule of each side. The anterior wall and partly the roof and

Endoskeleton of Frog

floor of each capsule are fonned by an irregular cartilage bone, the prootic. It is partially covered on its outer dorsal surface by the squamosal (Fig. 3A). Supraoccipital and basioccipital are absent. Dorsal side of occipital region is covered by the great frontoparietals, while its floor is occupied by the dagger-shaped parasphenoid. 2. Sphenethmoid. It is a tubular bone encircling the anterior end of cranial cavity, and fonning the posterior wall of nasal cavity. It is divided by a transverse partition into an anterior ethmoidal region and a posterior sphenoidal region. The latter encloses the fore-brain while the fonner is further divided by a longitudinal partition into right and left portions, each enclosing an olfactory sac. The bone is exposed only on its two lateral sides, as it remains covered ventrally by the parasphenoid and dorsally by the two nasals and fronto-parietals except for a small diamond-shaped area (Fig. 3B). 3. Fronto-parietals. These are a pair of long, broad, flattened and membrane bones. They are united along the mid-dorsal line and fonn the whole roof of cranium. In larval frog, each fronto-parietal occurs into separate frontal and parietal parts, but in adult frog, they become fused to fonn a single fronto-parietal. They extend in front overlapping the sphenethmoid and behind upto the exoccipitals. Anteriorly the two bones slightly diverge so as to expose a small diamond-shaped area of the sphenethmoid. Anteriorly they articulate with the nasals while posteriorly with the prootics and the exoccipitaIs (Fig. 3C). 4. Parasphenoid. The entire floor of cranium is covered and strengthened by a large parashpenoid bone which appears like a dagger or .1 in shape. The blade, shaft or pointed long arm of the dagger is directed anteriorly below the sphenethmoid while its handle with the cross-piece lies across the auditory capsules (Fig. 3D).

[II] Sense capsules There are a pair of auditory cap.l/t/es, enclosing the organs of hearing; and a paIr of olfactory capsules lodging the organs of smell, which are finnly fused with the cranium. They are mostly

[ 559 cartilaginous. The optic capsules enclosing the eyes are not fused with the skull. 1. Nasals. The anterior dorsal region of skull carries a pair of large, flat and triangular membrane bones, the nasals, which fonn the roof of olfactory capsules. The two nasals lie in contact in the median line. Their anterior ends reach up to the dorsal processes of premaxillae and partly fonn the boundry of the external nares. Their outer processes meet with the maxillae. Their posterior processes slightly diverge, before meeting the fronto-parietals, so that a small' diamond-shaped gap is enclosed where the sphenethmoid is visible dorsally (Fig. 3F). 2. Septomaxillaries. A minute irregular-shaped bone, the septomaxillary, lies close to the anterior process of each nasal. It is fonned by a broad basal plate and a pair of backwardly directed processes (Fig. 3E). 3. Vomers. Ventrally the floor of olfactory capsules is covered by a pair of ~mall flat, membrane bones, the vomers. Irregular or somewhat triangular in outline, they fonn the inner margins of posterior nostrils. The postero-lateral edge of each vomer bears a few pointed vomerine teeth in a row (Fig. 3G).

[III] Visceral skeleton The upper and lower jaws, the hyoid apparatus, the columella auris and the cartilages of the larynx, constitute the visceral skeleton. 1. Upper jaw. Premaxillae. The anterior-most bone of the maxiIIary arch or upper jaw on each side is premaxilla. It is a small irregular bone in the anterior tip of the snout, meeting its fellow in the middle line. Each premaxilla bears a few conical teeth in two rows along its anterior lower edge. Dorsally, it gives off a posteriorly directed process which fonns part of the inner boundary of external nostril. On outer side the premaxilla meets the maxilla of its side (Fig. 4). Maxillae. Each premaxilla articulates behind with a maxilla which fonns the greater portion of the outer margin of upper jaw. It is a long, thin and slightly curved bone provided along its whole length with numerous sharp, pointed and back(2-3)

Endoskeleton of Frog

560 J

~eriOrlimb

auditory capsule

premaxillary teeth

~il}ner' 11mb

maxillary teeth fenestra (Outer view)

o~osterior ~

(Dorsal view) PREMAXILLA

limb

(Inner view) PROOTIC

(MAXILLA)

PTERYGOID

Fig. 4. Frog. Loose skull bones of upper jaw.

wardly directed conical teeth, firmly ankylosed with the bone. Maxilla is connected behind with quadratojugal while it articulates at about the middle of its length with palatine and anterior limb of pterygoid. Quadratojugals. Quadratojugal is a small, slender bone with a characteristic comma-like shape. It lies behind L.1axilla forming the posterior part of outer margin of upper jaw. Its posterior broad end unites with quadrate cartilage. Quadrate cartilages. The suspensorium at each angle of the mouth, connecting the mandible with the skull, is formed by a small thin rod of cartilage, named the quadrate cartilage. In adult frog, it is rather completely covered over by the pterygoid and squamosal. The inner or anterior end of quadrate cartilage is fused with auditory capsule and outer or posterior end attached to the hind end of quadratojugal. Squamosals. Squamosals arc T-shaped or hammer-shaped bones attached dorso-Iaterally to the posterior end of cranium above the pterygoids, and helping to support the annular tympanic bones. The anterior limb of T or head remains free while the shorter posterior limh is attached to auditory capsule and prootic. Stem or handle of T articulates with the quadrate cartilage. Squamosal also forms the posterodorsal margin of orbit of its side. Pterygoids. These are present ventral to squamosals attached laterally to posterior end of cranium. Each pterygoid is a large three-rayed or Y-shaped bone. Its anterior ray or limb joins with maxilla and outer end of palatine. Its inner ray joins with parasphenoid and auditory capsule. Its (Z-3)

posterior ray articulates with quadratojugal and quadrate cartilage. Pterygoid contributes to the postero-ventral margin of orbit of Its side. Palalines. The ventral, anterior side of each orbit is bounded by a transver~ely elongated, slender, delicate, rod-like bone, called palatine. It connects the anterior side of cranium with the middle of maxilla. 2. Lower jaw or mandible. Lower jaw or mandible, is semicircular in outline. It is composed of two halves or rami, united anteriorly by a ligament. Teeth are altogether absent. Each half or ramus consists of a core of Meckel's cartilage surrounded by three bones, as follows (Fig. 5): Mento-meckelian. It is a small cartilage bone formed as an ossification at the extreme anterior end of Meckel's cartilage, at the anterior symphysis of two halves of mandihle. Angulosplenial. It is a long and curved bone forming most of the inner and posterior portion of each ramus of mandible. Its anterior end is tapering while its posterior extremity bears dorsally a condyle or knob-like articular surface for the mentomeckelian

coronary process

articular facet

~,.~

denta~

ANGULOSPLENIAL

~

angulosplenial

DENTARY (Inner view)

coronary process articular facet for quadrate

(ou~r view)

~' \

COMPLETE MANDIBLE

'\.) DENTARY (Outer view)

Fig. 5 Frog. Bones of mandible.

Endoskeleton of Frog quadrate cartilage of the skull. Just in front of this articular surface, it is produced upwards into a small coronary process. Dentary. It is a small, flat, dagger-like -bone, covering the outer surface of anterior portion of Meckel's cartilage. It extends anteriorly up to Mentomeckelian bOlle, whereas its posterior part runs closely applied to outer side of angulosplenial. 3. Hyoid apparatus. In tadpole, the skeletal framework supporting gills is formed by hyoidean arch and branchial arches. In adult frog, gills disappear and their skeletal framework is also reduced to form hyoid apparatus. It lies below tongue in the floor of mouth and provides surface of attachment to the tongue (Fig. 6). Hyoid apparatus is almost exclusively cartilaginous and is made of three parts : Body of hyoid. It is a thin membranous and broad squarish plate of cartilage, formed by the fusion of ventral ends of visceral arches. It lies beneath the tongue and gives off several processes. A short process is found at each angle of the plate. Anterior cornua. Anteriorly, body of hyoid gives rise to a pair of long, slender and curved, rod-like cartilaginous processes, called anterior cornua or horns. These are homologous with ceratohyals of lower forms. They run anteriorly, then curve backward and finally upward to become attached with auditory capsules of skull just below fenestra ovalis. Posterior cornua. Posteriorly, the body of hyoid gives off a pair of short but stout, straight and diverging ossified (bony) processes, called posterior cornua or thyrohyals, corresponding to the fourth branchial arches. They run one on either side of glottis or arytenoid cartilages supporting the laryngo-tracheal chamBer. 4. Columella auris. In the living frog, the cavity of each middle ear contains a single rod-like ear ossicle, called columella auris. It is made up partly of bone and partly of cartilage. At its outer end, it is attached to the middle of tympanic membrane or eardrum. Its inner end is inserted upon the outer wall of auditory capsule in

[ 561 anterior cornu

~alar

process

ntbOdY of hyoid

ClL

,.,..--~-,

posterior lateral

~ocess

posterior cornu Fig. 6. Frog. Hyoid apparatus.

m~~:~~

inner end of columella attached to fenestra ovalis

COLUMELLA AURIS

attached to tympanic membrane

Fig. 7. Frog. Columella auris.

an aperture called fenestra ova lis. Columella serves to conduct sound vibrations from tympanic membrane to the inner ear located deep in the head (Fig. 7).

Vertebral Column Vertebral column lies mid-dorsally In body, hence also called the backbone. It encloses and protects the spinal cord. Vertebral column of frog is exceptionally short in the absence of a tail. It is formed by ten separate bony elements arranged one behind the other in a linear series. First nine bones are small, ring-like and termed vertebrae. The last or the 10th bone is long, rod-like and termed urostyle (Fig. 8). 1. Atlas vertebra. The first vertebra is called atlas. It is in the form of a very small bony ring with reduced centrum and neural spine and without transverse processes and prezygapophyses. Anterior face of centrum carries a pair of large concave facets into which, fit the knob-like occipital condyles of skull. Postzygapophyses are present on the posterio( margin of neural arch. I

'(Z-3)

Endoskeleton of Frog

562 J facets for occipital condyle

neural spine /

transverse process

~~pre.zygp. post. zygp~.~~-neural arch posterior convexity of centrum

anterior concavity of centrum

TYPICAL VERTEBRA (Lateral view)

ATLAS (Dorsal view)

transverse process

4

'!-

neural spine centrum

posterior concavity of centrum

post. zygp.

n::~::;::~ ::a~.-/l ~~ ~~_~ ~ process

~

pre. zygp.

pre.zygp.

Or~

TYPICAL VERTE13RA (Anterior view)

posterior concavity of centrum

pos~ process

e~__J...... pre.

zygp.

anterior concavity of centrum

anterior concavity of centrum

SECOND (Anterior view) -

anterior concavity of centrum

TYPICAL VERTEBRA (Dorsal view)

TYPICAL VERTEBRA (Posterior view)

neural spine ~st.Zygp.

FOURTH (Antenor view)

EIGHTH (Ventral veiw)

neural spine

cavity for spinal cord concavities for 9th vertebra NINTH (Ventral view)

NINTH (Dorsal view)

UROSTYLE (Lateral view)

Fig. 8. Frog. Vertebrae.

2. Typical vertebra. In frog, third to seventh vertebrae are typical in structure. Each has a ring-like form with a large hole. called neural canal, through which the spinal cord passes. The ventral thick and solid rod-like part of the ring is centrum ,or body. It is procoelou.l' as its anterior (Z-3)

face is concave and posterior face convex. Dorsal arch of the ring is called neural arch. On either side, at its junction with the centrum, is given out an outwardly directed long and tapering process, called transverse process. Neural arch bears a small and blunt mid-dorsal process, the neural

Endoskeleton of Frog spine, which is obliquely directed backwards. Anterior margin of neural arch bears, at the base of neural spine, upwardly, and inwardly directed articular facets, called prezygapophyses. Posterior margin also bears similar pair of postzygapophyses which are directed downwards and outwards. 3. Second vertebra. Second vertebra is typical in structure with a slight variation. Its neural spine is short and conical while transver~e processes are small, and distally broad and flat. 4. Fourth vertebra. It is also typical in structure except that the transverse processes are broader distally. 5. Eighth vertebra. It resembles a typical vertebra In all respects, but Its centrum IS amphicoelous or biconcave. The anterior concavity receives the posterior convexity of seventh vertebra, while its posterior concavIty receives the anterior convexity of ninth vertebra. 6. Ninth vertebra. Ninth or sacral vertebra differs in many respects from the typical structure. Its centrum is biconvex, bearing one anterior convexity, fitting into the posterior concavity of eighth vertebra, and two posterior convexities fitting into the ant!!rior concavities of urostyle. Transverse processes are large, cylindrical, stout and directed backwards, their distal ends supporting the ilia bones of the pelvic girdle. Neural spine is inconspicuous. Pre-zygapophyses are well developed while Post-zygapophyses are entirely absent. 7. Urostyle. The posterior unsegmented part of vertebral column is called urostyle which is as long as the rest of vertebral column. It is somewhat triangular in outline with the pointed apex directed backward. Its centrum is rod-like with a broad anterior face bearing two concavities for articulation with the ninth vertebra. Its dorsal surface is raised up in the form of a prominent vertical ridge, highest in front but gradually tapering behind. Anteriorly, the ridge contains a short narrow neural canal which encloses the terminal part of spinal cord. Close to the anterior end, on either side opens a small aperture for the exit of tenth spinal nerves. These apertures correspond to the intervertebral foramina thus

{ 563

reflecting the compound nature of urostyle, but they are generally absent in Rana tigrina. Sternum. Sternum lies in the mid-ventral line intimately connected between the two halves of pectoral girdle. It includes four parts. Episternum is a flat circular and cartilaginous disc lying anteriormost. Omosternum is a bony rod connecting it with the clavicles. Mesosternum is a cartilaginous rod projecting behind the epicoracoids. Xiphisternum is the terminal broad, cartilaginous plate. Ribs are absent in frog.

Pectoral Girdle Pectoral girdle of frog is in the form of an inverted arch, buried in bodywall around the thoracic region of body. It protects the inner softer parts and provides support and attachment to the anterior limbs and their muscles. It is made both of bones and cartilages, and consists of two similar halves united midventrally with sternum but separated dorsally. Each half is further divisible into two regions (Fig. 9) : 1. Scapular region. It is the dorso-Iateral region comprising two bones. Suprascapula is a broad, flat, somewhat rectangular plate covering dorsally the first four vertebrae. Its upper free margin is made of calcified carti lage; while the lower part is bony and articulates with scapula. Scapula is a stout, flat bone, broader towards the ends but constricted in the middle. Posteriorly it forms the upper half of a deep cup-like depression, the glenoid cavity, for articulation with the head of humerus. A prominent conical process arises from its anterior side. clavicle

epicoracoid Fig. 9 Frog. Pectoral girdle and sternum In ventral view.

56,4

Endoskeleton of Frog

J

2. Coracoid region. It includes two bones and two cartilages. From the lower end of scapula run two bones, clavicle and coracoid, to unite mid-ventrally with sternum and their fellows of other side through a strip of cartilage, called epicoracoid. The clavicle is a slender rod, separated from the coracoid by a wide gap or coracoid foramen. A narrow stnp of calcified cartilage, called precoracoid, lies attached to it posteriorly. The coracoid is dumb-hell shaped with its inner end broader than the outer one which forms the lower half of the glenoid cavity.

facet for 9th vertebra ilium iliac crest articular L~~-:-- surface for urostyle

Pelvic Girdle Pelvic girdle of frog is a V-shaped structure occupying the posterior region of trunk. It gives support to pelvic region and hind limbs. Two limbs of the V run parallel to vertebral column. They diverge anteriorly but converge posteriorly and unite into a median disc which supports the posterior end of urostyle. Each lateral side of the disc carries a prominent cup-like depression, the acetabulum, for articulation with the head of femur. The girdle is made of two similar halves. Each half, or os innominatum, includes three elements all of which share the disc and the acetabulum (Fig. 10). 1. Dium. It forms the major part of the os innominatum. It is a greatly elongated bone running forwards to meet the transverse process of ninth vertebra. It is dorsally produced int!) a prolllinerit vertical ridge, the iliac crest. The two ilia meet posteriorly at an iliac symphysis in the median plane and form the anterior and upper half of the disc and acetabulum. 2. Pubis. Pubis is much reduced, triangular piece of calcified cartilage, occupying the ventral part of the disc and sharing nearly one-sixth of acetabulum. It is completely fused with its fellow of other side in a median pubic symphysis. 3. Ischium. It forms the posterior one-third of the disc and acetabulum and unites completely with the bone of the other half at a median ischiatic sy.mphysis. It is slightly bigger than pubis and oval in shape.

A

Fig 10. Frog. Pelvic girdle. A-Dorsal view. B-One half (os innominatum) in lateral view.

Forelimb Bones 1. Humerus. Bone of the upper arm is called humerus. It is a short and cylindrical bone with a slightly curved shaft. Its proximal end is swollen to form a rounded head, which fits into the glenoid cavity of pectoral girdle. Head is covered by calcified cartilage. Below the head, the proximal half of shaft bears anteroventrally a prominent vertical process, the deltoid ridge, to which muscles are attached. Distal end shows a round prominence, the capitulum or trochlea, with a condylar ridge on either side. It articulates with the groove of radio-ulna (Fig. 11). 2. Radio-ulna. Forel!f1ll contains a compound bone, called radia-ulna, formed by the fusion of radius and ulna bones. Proximally, the radio-ulna shows a concavity to receive the trochlea of humerus and further projected into an olecranon process forming the elbow joint. Distal half of

Endoskeleton of Frog

{ 565 head

olecranon process

distal epiphysis intermedium

deltoid ridge

radiale trapezium

articular facet for olecranon process of ulna

(Posterior view)

(Anterior view) RADIO-ULNA

HUMERUS

BONES OF HAND

Fig. II. Frog. Fore limb banes.

radio-ulna is somewhat flat and a groove imperfectly divides it into an anterior or radial part and a posterior or ulnar part, each terminating into a facet for articulation with the proximal row of carpal bones. 3. Bones of hand. Bones of wrist or capus are called carpals. In frog, carpal bones are six in number and arranged in two rows, each 'containing three. Bones of proximal row, which articulate with radio-ulna, are called radiale, intermedium or centrale and ulnare. Bones of distal row, which articulate with metacarpals, are termed trapezium, trapezoid and capitatohamatum. head

Hand or manus is supported by five slender, rod-like bones, the metacarpals. First metacarpal is rudimentary. Manus bears only four digits. Pollex or thumb is absent. Digits are internally supported by short bony rods, called phalanges. First and second digits bear 2 phalanges each, while third and fourth digits bear 3 phalanges each.

Hind limb Bones 1. Femur. Thigh is supported by a single long and slender bone, called femur. It has a slightly curved shaft and expanded ends covered by calcified

proximal articular facet

fibulare or calcaneum tiblale or

proximal epiphYSIS

astragalus~

tibial crest

:;:::::::::.§~-::::--2nd

polar body

clear cytoplasm

chromosomes of ovum and sperm mesodermal crescent Fig. I. Branchiostomll. Fertilized egg or zygote in dursal view.

animal pole. During fertilization, the sperm enters the egg near the vegetal pole (Fig. I).

?resumptive Area The fertilized egg or zygote is mosaic, diploid and has 24 chromosomes. It immediately gives out a second polar body and its cytoplasm becomes rearranged to form presumptive areas that form definite future organs of the animal. Clear cytoplasm of anterior half forms ectoderm, yolky

Development of Branchiostoma

fertilization membrane

[ 631

DORSAL

yolk cytoplasm (presumptive endoderm)

presumptive notochord

.b;~~:~;~:~me~

2nd polar body

POSTERIOR

fertilized egg

2-cell stage

2nd polar body clear cytoplasm (presumptive ectoderm)

4-cell stage

mlcromeres /~

chromosomes VENTRAL

Fig. 2. BranchirHtoma Presumptive arcas of zygote

In

side

VICW

cytoplasm develops into endoderm, whereas a crescent shaped granular area gives rise to mesoderm. At the animal pole, area of clear cytoplasm developed by rupture of germinal vesicle. Granular cytoplasm located in peripheral region flows downwardly towards vegetal pole and condenses to form gray crescent (Fig. 2).

Early Embryonic Development 1. Cleavage. Cleavage is quick process, starts soon after fertilizatio:1. Roughly it begins at sunset and completed upto morning. Before the outset of cleavage, vitelline membrane get separated from ooplasm and a wide cavity appears around ooplasm. Zygote of Branchiostoma contains very little amount of yolk so that its segmentation or division is total or complete, called holoblastic cleavage. The first and second cleavages are meridional or vertical and at right angles to each other resulting in 4 equal cells or blastomeres. Third cleavage is equatorial passing horizontally a little above the middle line producing 8 blastomeres of which upper 4 are smaller and called micromeres, and the lower 4 are larger called megameres. After sixth cleavage the embryo forms a ball-like mass of 64 cells, called morula (Fig. 3). 2. Blastula. Further cleavages result in a hollow ball, the blastula. It has a fluid-filled internal space, the segmentation cavity or blastocoel. Blastula is '1ot spherical but somewhat pear-shaped, the posterior end slightly pointed. Its wall is thin, only one-cell thick and the cells destined to produce specific structures. Smaller,

a-cell stage

16-cell stage

32-cell stage

Fig. 3. Brcmchiostoma. Cleavage upto 32-cell stage.

columnar micromeres on the ventral side form the future ectoderm. Larger yolky macromeres on the dorsal side form the future endoderm. Granular cells on its postero-Iateral sides are the potential mesodermal cells. A small antero-dorsal area contains small ectodermal potential notochord or DORSAL

ANTERIOR POSTERIOR

presumptive nervous system

future mesoderm

2nd polar body

future ectoderm VENTRAL

Fig. 4. Branchiostoma. Side view of complete blastula showing presumptive areas.

potential notochord

DORSAL

potential

potential nervous system

VENTRAL

potential ectoderm

Fig. 5 BrallL'iliostol/la. Right saglltal half of blastul.l.

Development of Branchiostoma

632] presumptive notochordal cells

mesoderm

neural plate

presumptive neural cells or nervous system

presumptive endodermal plate

ventral lip of blastopore endoderm

blastocoel disappearing

c

B

A

endoderm

Fig. 6. BranchlOstoma. Stages in gastrulation. A-Early gastrula. B-Mid gastrula. C-Completed gastrula.

chorda celis, and below it a small antero-ventral area includes ectodermal potential neural plate cells (Figs. 4 & 5). 3. Gastrula. The changes which convert the single-layered blastula into a two-layered gastrula, constitute gastrulation. It starts with the rapid proliferation of smaller ventral micromeres or ectodermal cells. As a result, the larger dorsal yolky megameres or endodermal cells first become flattened, then concave and gradually pushed into the blastocoel. This process is called invagination or emboly. It proceeds till the original blastocoel is completely obliterated and a secondary cavity, called archenteron, forms which opens to the exterior through a wide opening, the blastopore. The embryo now becomes bowl or cup-shaped and gastrula. Simultaneously, the double-layered presumptive notochordal cells roll inwards along

the dorsal edge or lip of blastopore followed by the presumptive mesodermal cells roIling inwards along the lateral and ventral lips of blastopore. This process is called involution. At the same time, the lips of blastopore grow backwards so that the larva elongates along its antero-posterior axis with a flat dorsal surface. The dorsal lip grows faster so that blastopore shi fts from dorsal to posterior end, becomes reduced to a small opening and persists as anus. The ectodermal cells forming the outer surface acquire cilia which help in the rotation of embryo inside its vitelline or fertilization membrane (Fig. 6). Organogenesis

Synchronized with gastrulation are the process of formation of neural tube (neuralization), formation of notochord (notogenesis) , and formation of neural groove

neural plate

A

overlapping ectoderm or neural fold

archenteron

B

c

gut wall (endoderm)

D

Fig. 7. Branchiostoma. Successive stages In the formation of central nervous system, notochord and mesodermal sacs, as seen in transverse sections. A-Completed gastrula B-Early neurula C-Mld neurula. D-Late neurula.

Development of Branchiostoma mesoderm (mesogenesis) and coelom. For convenience of study, the formation of each organ will be described separately (Fig. 7). 1. Formation of neural tube. The mid-dorsal ectoderm cells of gastrula become thick and large to form a neural plate. The centre of the plate is depressed to form a neural groove. Due to faster growth the cells along its two sides rise up to form longitudinal ridges called neural folds. The two folds meet each other and fuse over the groove forming a closed longitudinal hollow neural tube. The cavity of neural tube, or neurocoel, anteriorly opens to outside by a minute aperture, the neuropore, which closes in the adult forming the olfactory or Kolliker's pit. Posteriorly, the neurocoel encloses the blastopore and opens into archenteron by a small neurenteric canal which closes somewhat later. 2. Formation of notochord. The chorda cells situated middorsally in the roof of archenteron form the notochordal plate. It bulges up, towards the neural plate and finally pinched off from the archenteron to become the notochord. It elongates with the embryo, forming a solid mid-dorsal skeletal rod consisting of a linear row of vacuolated cells. Afterwards, it is surrounded by a notochordal sheath and also extends into the rostrum. 3. Formation of mesoderm, myotomes and coelom. At the time of notogenesis, the presumptive mesodermal cells in the dorso-lateral roof of archenteron also become deeply folded nerve cord

[ 633

neural folds forming central nervous system neural groove

~;:--~

_---::~~~~~~~~ pouches

notochord gut wall (endoderm)~~~[1

~~Jfi""'''''''cIDell[)mir. cavity

~~~~

Fig. 8. Branchiostoma. Stereogram of a post-gastrular stage to show arrangement of mesodermal pouches.

forming a longitudinal fold or groove on either side. Transverse partitions appear to divide each mesodermal groove into a linear series of mesodermal or coelomic pouches which are in continuation with the archenteron. The mesodermal pouches represent the beginning of metameric segmentation in Branchiostoma. The pouches subsequently cut off from the archenteron forming closed mesodermal sacs or coelomic cavities. Thus the coelom is enterocoelic in origin. After separation of notochord and coelomic sacs, the archenteron is reduced to enteron and develops the midgut diverticulum of the anteriormost or first pair of sacs, the left one remains small and opens into the oral hood forming the Hatschek's pit and wheel organ. The right sac enlarges to form the dorsal finray box mesodermal segment or somite dorsal aorta horizontal partition epidermis lateral plate mesoderm gut wall

A

of pouch

sublntestinal vein

Fig. 9. Brallc!lwstolllll. Further stages in T.S. to show development of mesodermal SOllutes and coelom

634

J

Development of Branchiostoma

neuropore

coelomic cavity

new somites forming from mesoderm ridge \

neurentenc

=~~nal notochord anterior gut diverticula

-)~ archenteron with evaginating somltes

mesoderm strand

Fig. 10. BnlllchlOstoma. Late neurula.

cavity of rostrum in front of the myotomes. The other mesodermal sacs on either side become each segmented into a small dorsal somite and a large lateral plate mesoderm. The dorsal somites remain distinct as myotomes and their cavities are called myocoels. The lateral plate mesoderms, flanking the enteron, grow ventrally pushing between the ectoderm (bodywall) and the endoderm (gut), and meet underneath the gut forming a temporary ventral mesentery. The cavity of each lateral mesodermal sac is called splanchnocoel. Its outer wall, attached to body wall is called somatic mesoderm, while the inner wall attached to the gut is called splanchnic mesoderm. With the rupture of partitions and ventral mesentery, all the cavities of the right and left sides become confluent forming a perivisceral coelom continuous underneath the gut (Figs. 8 & 0).

Neurula The embryonic stage is now termed a neurula. An identical neurala is formed in all vertebrates although its manner of attainment differs widely (Figs. 10 & 11). The neurula ha~ the following typical parts : (1) Booly elongated, fish-like. (2) Outer single layered ciliated ectoderm. (3) Endoderm forming an enteron or gut. (4) Rod-like skeletal notochord above gut. (5) Dorsal hollow neural tube above notochord. (6) (7)

Segmented dorso-lateral mesodermal somites or myotomes. Pervisceral coelom, lined by somatic and splanchnic layers of mesoderm.

Fig II Bnlllcillostoma. Neurula in v'L.S.

Hatching The neurula hatches out by rupturing its fertilization (vitelline) membrane when only two pairs of mesodermal somites have formed. Its surface is ciliated and it swims in the surface water layers of the sea as a free-swimming larva. As the newly hatched larva has no mouth and anus, it can not feed at once.

Larval Development The larva elongates rapidly, becomes laterally compressed and pointed at both ends. Mouth appears as a circular aperture to the left of mid-ventral line and bears cilia. Neurenteric canal clo~es and anus develops mid-ventrally near the hind end. Tail forms behind the anus. First pair of pharyngeal gill slits form ventrally but soon shift to the nght side. The larva having eight pairs of gill slits remains unchanged for a long time and later become fourteen pairs. Gill slits open directly to outside through ectoderm. Endostyle forms gradually on the floor of pharynx. The larva starts feeding on planktonic food by ciliary method. A Two larval club-shaped organ develops. longitudinal ventro-lateral metapleural folds appear. On their inner sides grow horizontally epipleures which meet and fuse together ventrally enclosing a large atrial cavity (Fig. 12). It gradually reduces the coelomic cavity and opens to outside though atriopore. The gill-slits now open into the atrium instead of opening directly to the exterior. The

[ 635

Development of Branchiostoma dorsal finray box

subatrial ridge

A

dorsal finray box

metapleural fold

c

B

Fig. 12. Branchiostoma. Three stages in T.S. showing successive development of atriulJI.

larva of Branchiostoma exhibit asymmetry unlike adult in the arrangement of various body organs like (1) Mouth appears first on left side. (2) Diverticula which originates from the anterior part of the gut also shows asymmetry. The left diverticulum remains smaller. (3) Development of gill slits also show asymmetry. Both set of gills first appear on right side.

Metamorphosis The larva, after passing three months pelagic life, now sinks down to the bottom, takes a burrowing life and gradually metamorphoses into the adult in several years. During metamorphosis, ectodermal cilia and larval club-shaped organs disappear and mouth becomes anterior. Oral hood with cirri, wheel organ and velum develop. Number of gill-slits increases due to subdivision of primary gill slits by tongue bars. Nephridia form, tail region elongates and notochord extends into rostrum. Sooner, the metamerically situated gonads develop and the adult lancelet starts breeding.

Significance

Study of the development of Branchiostoma IS significant from following point of view : (I) Study of embryology offers an insight into evolutionary history of chordates. (2) Position and way of the formation of blastopore in developing embryos divide bilaterally symmetrical animal into 2 groups viz., Protostomia (where blastopore marks the area of mouth) and Deuterostomia (where blastopore marks the anus). The Branchiostoma falls in the later together with other chordates and Echinodermates. (3) Way of formation of coelome in Amphioxus brings it closer to Echinodermates. Based on fate of blastopore, manner of mesoderm formation, muscle chemistry and similarity in sera proteins, it is believed that Coelenterates onwards two stock of invertebrates evolved. Branchiostoma, supposed to be a representative of primitive chordate and links chordates with invertebrates and shows the evolutionary steps followed by vertebrates.

Development of Branchiostoma

636 ]

IMPORTANT QUESTIONS

»

Long Answer Type Questions 1. 2. 3. 4.

5.

»

Shori Answer Type Questions l.

» l.

2.

3.

4.

Give a detailed account of development of Amphioxus. Describe the development of Branchiostoma upto the fonnation of three genninal layers. Describe the development of coelom in Amphioxus and show how it is affected by the developing atrium. Describe the structure of larva of Amphioxus and its metamorphosis to reach the adult stage. Draw diagrams of T.S. of the larval Amphioxus showing development of atrium.

Write short notes on - (i) Cleavage in Amphioxus, (ii) Gastrula in Amphioxus, (iii) Organogenesis in Amphioxus, (iv) Orgenogenesis in Amphioxus, (v) Fonnation of mesodenn and coelome in Amphioxus.

Multiple Choice Questions Development of Amphioxus resembles with those of : (a) Coelenterates (b) Annehds (c) Echinodermates (d) All these Number of chromosomes in the fertilized eggs Branchiostoma are : (a) 24 (b) 12 (c) 20 (d) 21 Yolky cytoplasm in Amphioxus develops into : (a) Ectodenn (b) Endodenn (c) Mesoderm (d) Both b and c Cleavage in Amphioxus is : (a) Holoblastic (b) Meroblastic (c) Superficial (d) DiSCOidal

5.

of

6.

7.

Emboly occurs in Amphioxus development during : (a) Cleavage (b) Blastulation (c) Organogenesis (d) Gastrulation Which amongst the following is olfactory pit in Amphioxus : (a) Hatschek's pit (b) Kolliker's pit (d) Hasel's pit (c) Larval pit At the time of hatching of Amphioxus larva the number of formed mesodennal somites are : (a) 5 pairs (b) 4 pairs (c) 2 pairs (d) one pair

ANSWERS 1. (c) 2. (a) 3. (b) 4. (a) 5. (d) 6. (b) 7. (c)

Development of Frog Frogs lay their eggs in water in early spring. During copulation or mating, the male firmly clasps the body of the female by his forelegs and enlarged thumb pads (nuptial pads). Smaller thickenings are also present below the joints of all digits of hand and foot, which are called articular pads. These also help in clasping the body of female. This sexual embrace is called amplexus, as already described in chapter 19. As the eggs are extruded through the cloaca of female (oviposition), the male deposits sperm cells over them (insemination). Thus fertilization is external, taking place in water. Each female lays several hundred eggs in a mass called spawn. The jelly that surrounds and protects the eggs, soon swells up by absorbing water, causing the eggs to stick to one another, so that the entire mass of spawn is held together. Development is indirect. the zygote forming a well-known aquatic larval form, called tadpole, which undergoes metamorphosis to become the terrestrial adult frog (Fig. 1).

Fig. I. Frog. Spawn.

Process of Fertilization Eggs are laid in the form of secondary oocytes, each containing the 1st polar body beneath the vitelline membrane near animal pole. Fertilization takes place before the jelly of egg swells up in water. During fertilization, the sperm always enters the animal hemisphere of egg just above the

Development of Frog

6381 animal pole

polar bodies

animal pole

vitelline or fertilization membrane pigmented animal hemisphere

coats of jelly (albumen)

previtelilne space

vitelline or fertllzation membrane

gray crescent

previtelline space vegetative pole vegetative pole

yolky vegetal hemisphere

Fig 3. Frog. Zygote withoutjdly in lateral vicw.

path. This dark streak is formed because some pigment granules from surface are carried deeper into cytoplasm along with the sperm head. The male and female pronuclei are haploid. Their fusion results in the formation of zygote nucleus restoring diploid chromosome number which is 26 in frog (Figs. 2-4).

Fig. 2. Frog. Fertilized ovum or zygote in dorsal view.

equator, losing its tail in the process. At the point of entry, the egg surface is raised into a small papilla or the cone of reception. After entry of sperm, some cortical reaction takes place due to which vitelline membrane get separated from the plasma membrane of the egg and a space called perivitelline space is established between the two. This space is subsequently filled with a liquid substance released by exocytosis of cortical granules which finally adhere to the inner surface of vitelline membrane. The vitelline membrane of egg becomes the fertilization membrane. Wolf et al., (1976) was of opinion that this membrane checks the polyspermy. Passage of sperm head (now male pronucleus) towards egg nucleus (now female pronucleus) is marked by a dark streak, consisting of a preliminary straight penetration or sperm path and a subsequently inclined copulation

Effects of Fertilization Entry of sperm brings about rearrangement of cytoplasmic materials of egg. The following changes are visible : (1) Even before the actual fusion of male and female pronuclei, the second maturation division of ovum occurs with the formation of a second polar body. (2) Cytoplasm of zygote contracts, expelling a small quantity of fluid. As a result, the vitelline membrane separates from egg surface, becomes more obvious and known as polar

1st polar body

vegetative pole

A

yolky vegetal hemisphere

female pronucleus (haploid)

B Fig. 4. Frog. Stages in fertilization of ovum.

c

zygote

[ 639

Development of Frog animal pole (dorsal)

1st cleavage

blastomeres

smaller micromeres

2nd cleavage

o segmentation cavity

micromeres

micromeres

blastocoel

(yolk cells)

E

F

G

H

Fig. 5. Frog. Cleavage and blastulation. A-Zygote. B-2-cell stage. C-4-cell stage. D-S-cell stage. E-Early blastula. F-V.S. early blastula. G-Late blastula. H-V.S.late blastula.

fertilization membrane. This also enables the egg to rotate freely. (3) In the side opposite that of spenn entry, there is an inward influx or movement of superficial pigment. This results in the fonnation of a crescentic surface area in the animal hemisphere near the equator. As this region becomes intennediate or grayish in pigmentation instead of the nonnal dark, it is known as the grey crescent which fonns an important landmark. It marks the position of dorsal lip of future blastopore and the region from which mesoderm and notochord will later develop. Before fertilization, the egg was radially symmetrical. With the appearance of grey crescent, it becomes bilaterally symmetrical. With respect to the future embryo and adult, the grey crescent marks the future dorsal surface, the animal pole marks the anterior end, and the vegetative pole the posterior end.

Early Embryonic Development Cleavage Jelly present around the egg imbibes water and swells up as a result of which eggs get separated from each other which provides favourable environment for eggs. All the subsequent developments and divisions take place within this vitelline membrane and jelly. Fertilization IS followed by cleavage or segmentation of zygote, which is holoblastic but unequal due to large quantity of yolk present. First cleavage is meridional, passing vertically from animal to vegetative pole. It divides the grey crescent and zygote into two similar and symmetrical cells, called blastomeres, which represent the right and left sides of future embryo and adult. Second cleavage starts at the time when· the first cleavage furrow is in process of cleaving the yolky cytoplasm of the vegetal hemisphere is also

(Z-3)

640

J

meridional (vertical), passing through the poles, but at right angles to the first cleavage plane, producing four equal blastomeres. Third cleavage is latitudinal (horizontal) but well above the equator toward the animal pole, producing eight blastomeres of two distinct types. The upper four smaller pigmented cells at the animal pole are called micromeres. The lower four larger yolk-containing cells at vegetative pole are called megameres or macromeres. Further cleavages become less regular and difficult to follow. However, the smaller micromeres in the animal hemisphere divide much faster than the larger macromeres because of large quantity of non-living inert yolk present in the vegetative hemisphere (Fig. 5). Blastula formation A solid ball-like morula stage does not occur in frog. Instead, a hollow ball-like blastula stage is formed. As early as the 8-cell stage, a small central fluid-filled space, called segmentation cavity or blastocoel, appears within the embryo which is now termed a blastula. The blastocoel probably serves two major functions- firstly it is the cavity that permits migration of cells during gastrulation, secondly it prevents interaction of cells of vegetal half with animal half. In a fully formed blastula, blastocoel is a large hemispherical cavity entirely in the u{lper or animal half. Its dome-like roof is formed by numerous small, pigmented, black micromeres, while its floor is composed of large, yolk-laden, white macromeres. The blastula wall, or blastoderm, is one-cell thick in Branchiostoma, but many cells thick in frog. Gastrulation A series of chanees, converting the single-layered blas~l1la into a two-layered embryo or gastrula, are collectively known as gastrulation. It involves mass migrations (formative movements) and rearrangement of cells (or presumptive areas) of blastula. Several changes or processes taking place simultaneously during gastrulation are as described below. 1. Epiboly. A fold of rapidly dividing pigmented cells of micromeres of animal pole (Z-3)

Development of Frog presumptive notochord neural plate or presumptive nervous tissue

presumptive epidermis presumptive endoderm

dorsal lip 01 blastopore

presumptive endoderm

Fig. 6. Frog. PresumptIve areas on zygote or blastula. A-Postero-dorsal view. B-Side view.

gradually grows over the light-coloured yolky cells or megameres of vegetative pole. It completely encloses the megameres except in the region of yolk plug. This process of overgrowth is called epiboly. 2. Formation of blastopore. In the beginning of epiboly, a small crescentic groove appears postero-dorsally on blastula a little behind the edge of grey crescent in the presumptive endoderm. Its anterior pigmented margin is called the dorsal lip of blastopore. Its backwardly projecting lateral horns are called the lateral lips. As epiboly progresses, the lateral lips finally meet below forming the ventral lip. Thus, the crescentic groove becomes a complete circle, or blastopore, through which is visible a tiny white spot of yolky endodermal cells, called yolk plug. It is present ventrally slightly in front of vegetative pole. 3. Invagination of endoderm. As the prospective ectoderm cells or micromeres advance, the future endoderm cells or megameres gradually migrate towards blastopore and gradually sink inside. With the completion of blastopore, the whole of yolky megameres or future endoderm becomes internal. 4. Formation of archenteron. With the inturning of tissues the earlier crescentic groove gradually grows inward into a new cavity, the' archenteron or primitive gut, which opens to outside through blastopore. As a result of inwardly rotating endodermal cells, the blastocoel becomes gradually reduced and finally obliterated.

Development of Frog

[ 641 micromeres remains of blastocoel

archenteron

~~~~_YOlk plug

~

A

megameres (endoderm)

mesoderm

mesendoderm

B

Fig. 7. Frog. Stages of gastrulation in median sagittal sections. A-Early gastrula. B-Mid gastrula. C-Completed gastrula (yolk-plug stage).

5. Involution. In the beginning of epiboly, the advancing future notochord cells. or chorda cells. become inflected or turned inside round the dorsal lip of blastopore. They extend beneath the neural plate cells which however remain on the surface. With the formation of lateral lips of blastopore. the future mesoderm cells also roll inside over the lateral lips. Internally they occupy positions on either side of chorda cells between the surface epidermis and endoderm. Mesodenn of ventral side also rolls over the ventral lip of blastopore. In a completed gastrula. blastocoel is obliterated. Archenteron is well developed and opens to outside through blastopore later reduced to a slit (proctodaeum). Roof of archenteron is

made by notochord cells and sides of mesoderm cells. Floor of archenteron contains a mass of yolky endoderm cells visible through blastopore as yolk plug. The outer layer of pigmented cells consists of ectoderm and neural plate. Formation of gastrula is now complete. Fate maps in frog. The entire surface of the blastula may be divided into three regions(1) A large area on and around the animal pole. (2) Marginal zone which spreads all around the equator of the blastula. (3) The area on and around the vegetal pole. These areas can be differentiated on the basis of their pigmentation. Whole of the animal region is deeply pigmented. The marginal zone is much

lateral lip of blastopore

vegetal pole

A

megameres

yolk plug

B

c

o

ventral lip of blastopore

Fig. 8. Frog. Successive stages in the formation of blastopore as seen from vegetative pole (posterior view).

(Z-3)

Development of Frog

642 ] plate

mesoderm ectoderm

the notochord area the marginal zone areas are taken up by the material for the segmental muscles of the body. The lateral and ventral parts of the marginal zone give rise to the mesodermal lining of the body cavity, the kidney etc. The cells of the vegeted region take part in the formation of midgut and hindgut.

(yolk cells)

Formation of neurula stage (organogenesis)

archenteron Fig. 9. Frog. Gastrula in T.S. showing three primary germinal layers.

broader on one side of the embryo and is lightly pigmented, while the vegetal pole has very little pigmentation. Each of the above regions corresponds to the future organs of the animal. On the basis of the vital staining the cells of these areas can be differentiated which in future will form the various organs of the embryo. The animal region has two areas, one of which has to develop into the nervous system of the embryo and the other area becomes the skin of the embryo. The material for the sense organs is also present in these areas. Inside the nervous system area, a small area may be localized which takes part in the formation of the eyes of the embryo. Similarly in the epidermis area the material for the nose, ears and the foregur may be found. In the marginal zone the material for the notochord occupying a large area on the dorsal side of the blastula may be found. The area lying near the vegetal pole contains the material for prechordal connective tissue. Towards the vegeted pole but still inside the marginal zone lies the material for the alimentary canal, the endodermal lining of mouth, gill region and pharynx. On both sides of

Neurogenesis Towards the end of gastrulation, the presumptive neural plate area, running forwards from the blastopore, becomes thickened forming a mid-dorsal pear-shaped medullary or neural plate. Running throughout its length is an open midline depression or neural groove. This is the early neurula stage of embryo. The edges of lateral sides of neural plate rise up as a pair of ridges or neural folds. These folds grow to meet and fuse with each other in the midline so that the open neural groove is changed into a close neural tube, with its cavity called neurocoel. Its anterior wider end forms the brain and narrower posterior end the spinal cord. AnteriGr end opens outside for a short time by a neuropore. But it soon closes, and the anterior end of neural tube enlarges to form the primary cerebral vesicle, which is constricted to form fore-, mid- and hind brain vesicles. At the posterior end of embryo, the blastopore becomes drawn out as a longitudinal slit. Its lateral margins fuse in the middle of the slit forming the primitive streak and groove. However the blastoporal slit remains open at the upper and lower ends by neurenteric canal and proctodaeum. both opening ectoderm

ectoderm

neural plate

~~. ~.•'~ ~

.

(Z-3)

~.

neural groove

neural fold

~\/.~

neural)¢}~l?~"'" crest

neural crest

ganglion

neurocoel ]::::...."'---- neural tube

~"-':-~~

....

notochord~

@---- notochord ---~

C

Fig. 10. Frog. Stages in formation of neural tube (neurogenesis).

0

[ 643

Development of Frog into the archenteron. Posteriorly, the neural folds close over the neurenteric canal through which the neural tube opens into archenteron. Later, the neurenteric canal closes so that there is no connection between neural tube (spinal cord) and gut. The proctodaeum marks the site of future anus or cloaca. Certain ectoderm cells at the lateral margins of neural plate do not become incorporated in the neural tube. They remain separate forming linear bands called neural crests. They give rise to parts of autonomic nervous system, dorsal root ganglia of cranial and spinal nerves, and certain other tissues such as trabeculae of skull and elements of visceral arches (Fig. 10). Notogenesis

The chorda cells lying in mid-dorsal region of roof of archenteron, separate from adjacent mesoderm cells. They form a cylindrical rod-like notochord made of characteristic vacuolated cells. It becomes the axial skeleton of embryo. A notochordal sheath develops around the notochord. Formation of mesodermal somites

The presumptive mesoderm cells, inturning at the lateral margins of blastopore, become arranged on either side of notochord in two ~olid blocks or sheets without any cavity. This is unlike Branchiostoma in which mesoderm first appears as hollow mesodermal sacs. Mesodermal sheet on either side is differentiated into three fairly distinct zones. The most dorsal and thicker zone, close to notochord, is called epimere. The most lateral and thinner zone is hypomere or lateral plate mesoderm. The intermediate cell mass is called mesomere or nephrotome. The hypomere extends laterally and ventrally around the yolk cells, so that mesoderm forms a continuous layer beneath the archenteron which is now lined wholly by endoderm. Mesoderm now constitutes the third germ layer, lying between the other two, that is, endoderm and ectoderm. At this stage, the embryo becomes triploblastic. The epimere mesoderm of each side thickens and segmented transversely into blocks of cells,

the somites. Formation of somites begins in front of the middle of embryo and proceeds backwards. Nephrostome and lateral plate mesoderm remain undivided. Formation of coelom

A split occurs in the hypomere of lateral plate mesoderm separating an outer somatic or parietal layer and an inner splanchnic or visceral layer. The cavity thus formed between the two layers is splanchnocoel which marks the beginning of coelom. It extends downwards to become continuous with that of the outer side below get to appear U-shaped in a section. Thus coelom in frog IS a schizocoel formed by the splitting of mesoderm, in contrast to the enterocoel of Branchiostoma which is derived from the archenteron. Somatic lay~r unites with ectoderm to form the somatopleure or bodywall, while visceral layer unites with endoderm to form the gut wall or splanchnopleure. A narrow cavity, called myocoel, appears in each mesodermal somite which itself differentiates into 3 parts. Outer thin dermatome becomes the dermis of skin. Middle thick myotome forms the body muscles. Inner sclerotome contributes to vertebral portions of skeleton. Myocoels shortly disappear without contributing to the coelom of the adult. Myomeres lead to the formation of kidneys, and their cavities, the nephrocoels, persist as excretory tubules and their ducts which are therefore coelomic in origin. The splanchnocoles of hypomeres become the general body cavity or perivisceral coelom lint:d with peritoneum. Formation of eye

After the differentiation of the three primary brain regions from the neural tube, the rudiments of the eye start appearing in the region of future diencephalon. The presumptive retinal plate which at lateral sides of forebrain begin to evaginate as protsuding sacs. They are called primary optic vesicles. Each optic vesicle continues to grow towards the epidermis. By the pressure so developed the intervening mesenchyma is displaced and ultimately comes III contact with the

Development of Frog

644 ] surface ectoderm

~.

optic '"P

optic __ .' . :.. , '-'- - lens vesicle stalk __ .... : lens :.: ....... ::.

A

placodes

8

o

E

c

F

vitreous humour

Fig. II. Stages of development of eye of frog.

epidermis. The wall of the optic vesicle undergoes invagination to form a double walled optic cup-enclosing a space which is the future posterior chamber of the eye. The cup is two walled structure. Pigment granule.s develop in the external thin layer which forms the pigmented coat of retina. In the invaginated thicker layer sensory and conducting cells differentiate to form the neurosensory layer of retina. There two layers are continuous with one another at the rin of the optic cup which forms the edges of the pupil. In the beginning the pupil is quite wide but lateron its edges bend inward and converge so that the opening of the pupil is reduced. Edges surrounding the pipjl, become considerably thin to form the iris. Later ciliary body and retina is developed. Epidermis of the head opposite to the optic vesicle thickness ~o form the lens-placodes which invaginates to form the lens cup which gradually deepens and puiches off from the epidermis and fprms a closed lens vesicle. The lens vesicle gradually comes to lie in the secondary optic vesicle. Cells of the lens vesicle undergo

cytodifferentiation and ultimately develop into a refracting body (Fig. 11). Cornea is formed by the ski n epidermis' and mesenchyma of the head region. In larval stage the epidermis forms the external cornea which remains continuous with the skin while the mesenchyma forms the internal cornea which is continuous. During metamorphosis these layers fuse and become transparent. The choroid coat and the sclera of the eye develop from the mesenchyma cells accumulating around the eye-ball. The inner layer of mesenchyma gives rise to a network of blood vessels surrounding the pigmented retinal epithelium and is called choroid coat. The outer layer of the mesenchyma forms a fibrous capsule, the sclerotia coat or sclera around the eye which has protective function.

Formation of heart Embryonic development of the heart initiates from the lateral mesoderm plate in the pharyngeal region. After the formation of the nerve chord, the

Development of Frog

[ 645 endoderm

splanchnic layer of -""-'~~ mesoderm

8 pericardial

[email protected]

cardiac endothalial cells

C

somatic layer _.......;.=.of mesoderm pericardial cavity

splanchnic layer of mesoderm

dorsal mesocardium cardiac endothalial cells wall of gut

E

splanchnic layer of mesoderm

somatic layer of mesoderm

F

Fig. 12. Fonnation of heart in frog. A-B-Mesoderm reaching midline. C-Enlargement of pericardial cavity. D-Endothalial cells being arranged in the form of a tube. E-Heart tube established. F-Dorsal mesocardium stilJ present.

free edges of the mesodennal mantle gradually converge towards the mesodennal free area in the pharyngeal region. They get thickened in the region of the heart and thus form the heart rudiment. A number of loose cells are derived from the ventral edge of the mesodennal mantle. They form the endocardium. These cells accumulate in the middle line and get arranged to fonn the endocardial tube. This tube bifurcates at the two ends. At the anterior end it extends as aortae and

at the posterior end it receives two vitalline veins. Soon the lateral plate of mesodenn come closer and fuse with each other. The visceral layer of mesodenn envelops the endocardial tube on the dorsal side as well. By fusion of the mesodermal layers of the right and left sides, epithelial partitions are fonned above and below the endocardial tube and thus pericardial coelom is formed around the endocardial tube. The visceral layer of mesodenn gives rise to the myocardium.

646 J

Development of Frog

truncus arteriosus

ventricle dorsal rnesentry --::=2Z~~~~~~~~~5

ventricular portion _

~

sino-atrial ~ portion

.

} vitelline veins

- - - perlcardial cavity

bulbous ventral aorta arteriosus ~~ conus \ . ~\fo~/arteriosus

'-ctruncus

I

.~~ventriCle

'.'

"

~

c

TfbUlbOUS

vitelline ,veins

-:::======J~ ,,:::::L' "venosus sinus '-='--=------'--

jatriurn

ventricle

o

Fig, 13. Stages of development of heart ill frog,

The parietal layer of mesoderm forms the pericardium. The cells of the endocardial tube continue to mUltiply mitotically and as a result of this the tube undergoes a folding and attains an S-shaped structure. It becomes constricted in some places and dilated at others. In the posterior part where the vitalline views are present, the sinus venosus is formed. Anterior to sinus venosus the atrium is developed ventral to the atrium the thick walled ventricle is formed. Most anteriorly the ventricle forms two dilations called the conus anteriors and bulbus anteriosus. Later the single atrun gets divided into right and left atria. The two ventral aortae of arising from the ventricle give rise to the trunclls anteriosus (Figs. 12 & 13). Post-neural or pre-hatching embryo

Even during formation of neurula, the embryo begins to elongate and acquires a flat surface

eminence, called gill plate, on either side near the anterior end. Soon after completion of neurula, a postero-dorsal projection above the proctodaeal pit forms the tail bud. It grows out posteriorly into a post-anal tail. The proctodaeal pit breaks through into mesenteron to become the cloacal opening. Three ridges on either side on each gill plate mark the position of first three branchial arches. Behind them light dorso-Iateral elevations indicate the positions of first 2 to 3 pairs of mesodermal somites. On the ventral side of head is a shallow oral or ventral sucker containing mucous or cement glands. Anterior or above the sucker an oral depression, the stomodaeal pit, marks the beginning of mouth. While still within its egg membranes, the embryo moves about by means of cilia on its epidermis, with an occasional twitching caused by contraction of muscles developed from somites (Fig. 14).

Development of Frog neural plate

[ 647

neural folds

neural plate

neural folds

neural plate

neural groove

neural fold mesoderm primitive streak and groove lateral plate mesoderm

(utrue anus or proctodaeum

A rU?iments of

gO! ",ch"

endoderm (yolk cells)

c

B tail bud

~y~) I

spinal cord

, ;''''''£'''.. '. :...:~~myotomes ....... . .;;'>. '_'~~ /

,"

"

~

external naris

fornl;~: "~"\:;~;,;mb A

B

c

adult frog

Fig. 16. Frog. Stages in metamorphosis.

functions. It has a well-developed locomotory tail with caudal fin and muscles for swimming. It has eyes, external nostrils, mouth, long spirally coiled intestine, cloacal opening and spiracle which _are readily visible. Cement glands (ventral sucker) have disappeared. In respiration water passes through mouth, pharynx, branchial clefts, over internal gills into the opercular chamber and out again through the spiracle. In later stage lungs develop from pharynx so that tadpole uses both gills and lungs. Soon the branchial clefts close, internal gills absorbed and opercular cavity disappeared. The tadpole now frequently comes to the surface to take in air into buccal cavity and lungs for respiration. A well developed lateral line system is also present. Mesonephric kidneys are developed. Of the two pairs of limbs, the hind limbs appear first as tiny hemispherical elevations or buds at the base of tail. Buds of forelimbs are not visible at first being covered by the operculum. Later, the left forelimb emerges through the spiracle, while the right forelimb breaks through the wall of opercular pouch.

Metamorphosis Two or three weaks, after breathing with lungs, the tadpole undergoes drastic changes called metamorphosis, and converted into a tiny young frog differing from the adult only in size (Fig. 16). Some of the important changes are as follows ,: (1) Tadpole stops feeding but frequently visits water surface to engulf air into lungs via mouth and then sinks down to water bottom again. (2) Anterior limbs break through operculum. (3) Tail gradually shortens providing nourishment. (4) Head and body become increasingly frog-like. Eyes become prominent and legs become longer. (5) Ciliated larval skin, frilly lips and horny jaws are cast off. (6) Mouth grows wider, true jaws develop, tongue enlarges, stomach and liver also enlarge, but long and coiled intestine of predominantly herbivorous tadpole greatly shortens into small gut of carnivorous frog.

650

J

Development of Frog

(7) Bone formation begins in hitherto wholly

(8)

(9) (10) (11) (12)

»

IMPORTANT QUESTIONS

I.

2.

3.

4.

5.

6.

7

Describe Describe Describe What do Describe

the life history of Indian bull-frog, Rana tigrina. the development of frog upto the end of gastrulation. the development of frog upto fonnation of three genninal layers and mention their fate. you mean by metamorphosIs ? Explain with reference to the life history of frog. the structure of tadpole of frog. Discuss the metamorphic changes it undergoes to become the adult.

Short Answer Type Questions I.

»

to live in damp places on land. It feeds on insects and grows into the adult. Significance. Study of the embryology of frog is practically useful to us in variety of ways (1) It helps in interpretation of avian and mammalian development (2) It explains the evolutionary tranSItIOn of lower chordates into higher chordates. (3) It explains the evolution of lung breathing animals from gill breathing animals. (4) It also explains the evolution of various physiological requirements present in aIr breathing and land living animals.

Long Answer Type Questions I. 2. 3. 4. 5.

»

cartilaginous endoskeleton and musculature develops rapidly. Vascular system is modified for air breathing with lungs assuming more importance as respiratory organs. Skin becomes vascular, respiratory, pigmented and slimy. Pronephros IS replaced by mesonephric kidneys. Lateral line sense organs disappear. Young frog has a stumpy tail. It leaves water

Write short notes on- (i) Neurogenesis, (ii) Notogenesis, (iii) ~sumptive areas, (iv) Fate maps in frog, (v) Formation of heart in frog, (vi) Formation of eyes in frog, (vii) Formation of coelom and mesoderm.

Multiple Choice Questions On forelimbs of male frog, well developed pads are found for clasping the female at the time of copulation is called: (a) Nuptial pads (b) Tuberoclties (c) Suchial buds (d) None of these In case of frog, fertilization is : (a) Internal (b) External (c) Both these (d) None of these Development of frog is : (a) Direct (b) Indirect (d) None of these (c) Both of these Diploid number of chromosomes in the zygote of frog is : (a) 22 (b) 26 (c) 24 (d) 25 Fertilization membrane is formed in case of frog : (a) Before fertilizatton (b) After fertilization (c) After oviposition (d) Before oviposition First cleavage In frog is : (a) Holoblastic equal (b) Holohlasttc unequal (c) Meroblasttc (d) Hon711ntal Eggs of frog are .

8.

9.

10

II

12.

(b) Megalecithal (a) Mlcrolectthal (d) TeloleCIthal (c) Alecithal Blastula of frog IS called . (al Coeloblastula (b) Disl"Ohlastula (c) Holoblastula (d) Amphtblastula On the baSIS of pigmentatton surface of the blastula of frog can be divided mto : (b) Two regions (a) Three regIOns (c) Four regions (d) Five regions Coelom of frog is . (a) Acoelous (b) Pseudocoelous (c) Enterocoelous (d) Schizocoelous Coelom of frog is derived from: (a) Sphttmg of mesoderm (b) From archenteron (c) Both of these (d) None of these Innermost compartment of myocoel called Sclerotome contnbute to the formatton of : (a) DermiS (b) Muscles (c) Vertebral skeleton (d) All these

ANSWERS I (a) 2. (b) 3. (b) 4. (b) 5. (b) 6. (b) 7. (d) 8. (a) 9. (a) 10. (d) I \. (a) 12. (e).

Development of Chick (Fowl) It is preferable to study the embryology of chick or the common fowl (Gallus gallus) to that of pigeon (Columba livia) becau!>e of several advantages. Eggs of fowl are large in size, easily available throughout the year and can be incubated artificially. Moreover, the process of development has been most thoroughly worked out in fowl. A comparative study of embryology of different birds shows that it is essentially similar in all the birds with only minor unimportant differences. Embryology of birds is much like that of reptiles in general. Development is direct, without a larval stage.

Fertilization Ova leave the ovary (ovulation) as primary oocytes. They are released in coelom and caught by the expanded funnel-like opening of oviduct. They are fertilized in the upper part of oviduct which also receives sperms from the male bird during copulation. The sperm of cock have long lives about three weeks or so. Several (5 or 6) sperms enter the oocyte (polyspermy) which immediately undergoes two maturation divisions.

The resulting two polar bodies degenerate and disappear. The nucleus of only one sperm then fuses with the nucleus of ovum proper resultiQg in fertilization. The remaining sperms migrate to the periphery of the ovum and ultimately die. Thus fertilization is internal in birds. Fertilized ovum or zygote normally takes 24 .hours to pass ddwn the oviduct, before being laid. While descending, it is surrounded in succession by various envelopes of albumen, shell membranes and porous calcareous shell, all secreted by the wall of oviduct.

Structure of Egg of Hen or Fowl Shape and size. Generally the eggs are laid at the rate of one egg per day. A fully formed and newly laid egg is large and elliptical with one end broader than the other. It is about 3 cm broad and 5 cm long. By the time it is laid, the embryo is in the blastula stage or undergoing gastrulation. As the eggs are deposited outside water, on land, they have protective envelopes as a safeguard against drying and mechanical injuries. Egg shell. The egg is externally protected by a firm white or brown shell, at least 94% of

652

J

Development of Chick (Fowl) double

membrane

blastodisc (acrosome

b~OO"~

i:;.-- middle

~;:

middle layer of dense albumen

A

8

porous shell

c

yolk

outer layer of thin albumen

latebra

vitelline membrane

D

Fig. I. Fowl. A-Fowl 's sperm. B-Pigeon's sperm. C-Hen's egg partially in section. D-- V. S. ovum.

which is calcium carbonate. The shell is porous and allows diffusion of 02 and CO 2 through it. The shell is soft and flexible in a freshly-laid egg but soon becomes hard and brittle. The shelled-egg, shut off from its surrounding, is termed cleidoic (Fig. 1). Shell membrane. Immediately underneath the shell is a thin but tough, white shell membrane consisting mostly of keratin. It is made of two layers which are mostly in close contact with each other but are separated at the broad end of the egg to enclose an air space. As development proceeds, the air space grows larger, and a little before hatching the young chicken pierces the air space with its beak and takes its breath of air, inflating its lungs. Albumen. Beneath the shell membrane lies the albumen or white of egg, surrounding a central mass of yolk. The albumen consists chiefly of water (85%) and protein albumen. Other proteins are also present. The albumen is deposited in several layers. The outermost albumen is more water-like and known as the fluid, liquid, watery, or thin albumen. The middle layer of albumen is thick and viscous and known as the dense albumen. The innermost layer is made of very viscous albumen called the chalaziferous layer, which surrounds the yolk. It forms a pair of

spirally twisted ropes or cords, the chalazae, one towards each end of the egg. The way in which they are formed is uncertain, but probably they are produced by the rotation of the egg in the oviduct. They are formed by the fibres of a glycoprotein, called ovomucoid. Ovum. The true ovum or egg cell proper of fowl is very large, with its enormous mass of yolk. It is invested in a thin transparent vitelline or fertilization membrane, which separates the yolky ovum from the surrounding albumen. The membrane is formed by the union of a primary membrane with a secondary membrane deposited by the follicle cells of the ovary. The granules of yolk, being slightly denser than cytoplasm, sink to the bottom of the ovum, towards its vegetal pole. Due to the large yolk contents, the egg of fowl is an extreme example of polylecithal, macrolecithal and telolecithal egg. The yolk is not homogeneous in composition but consists of alternate layers of yellow and white yolk arranged concentrically around a central flask-shaped mass of white yolk. The layers of yellow yolk are thicker than those of the white yolk. Yolk consists largely of phospholipid lecithin and fat. Analysis of solid yolk shows 60% fat and 30% protein. The white yolk contains less fat and also less of the fat-soluable pigment carotene to which the yellow

[653

Development of Chick (Fowl) vitelline membrane

C

sub\lerminal space

G blastoderm

,

o

\'

yolk

E

Fig, 2, Fowl. Early stages in cleavage of the germinal disc, A-2-cell stage, B-4 cells. C-8 cells. D-16 cells. E-32 cells. F-154 cells. G-Y.S. of 32-cell stage, H-Hemisection of I 54-cell stage.

colour is due. The yolk contains about 50% water. The central flask-shaped mass of white yolk is termed latebra, which its outer neck-like part is known as the neck of latebra, which expands under the blastodisc into a broad disc, the nucleus of Pander. The ovum contains a nucleus surrounded by a negligible amount of yolk free cytoplasm forming the germinal disc or blastodisc. It always floats on the upper surface of yolk and represent the animal pole of the ovum. Yolk and albumen provide food (protein) to the developing embryo. Towards the end of incubation, a part of shell dissolves and the CaC03 released is utilized to build the developing bones. The shell also provides a little protein. The egg-white or albumen supplies water and minerals, while vitamins come from both yolk and albumen.

Cleavage or Segmentation Cleavage begins about 3 hours after fertilization As already stated, cleavage and early gastrulation are completed by the time the egg is laid. Cleavage furrows do not extend into the yolk but are confined to the tiny germinal disc (blastodisc) of cytoplasm occupying a small circular area on the top of yolk. First cleavage is a vertical furrow

between two daughter nuclei. It does not extend right across the disc, nor through its whole thickness. As a result the disc is divided only incompletely into two cells or blastomeres. Second cleavage furrow is also incomplete and at right angles to the first, resulting in four blastomeres. It is followed by more incomplete and irregular furrows in rapid succession. The entire blastodisc is thus cut up into a sheet of cells called the blastoderm. Since many of the cleavages are horizontal, the blastoderm is several cells thick. The partial and superficial cleavage in chick resulting in the formation of a blastoderm, is called discoidal and meroblastic in which yolk, fOlming the greater part of ovum, remains permanently undivided. It is different from the holoblastic cleavage of Branchiostoma and frog in which the entire ovum is involved in the division. A marginal zone of unsegmented cytoplasm, called periblast, unites blastodisc with the yolk mass, thus forming a zone of junction (Fig. 2).

Blastulation The free margin of blastoderm grows rapidly over the surface of yolk. Soon its central region IS separated from the underlying yolk by a

654

J

Development of Chick (Fowl)

marginal cells (area opaca)

ANTERIOR

area opaca ectoderm neural ectoderm

area pellucida

.....

notochordal mesoderm

"\

>-:.:

"

),,;~.

,~.

"

~~ r~

limit of embryonic epidermis

yolk mesoderm

POSTERIOR

A Fig. 3. Fowl. V. S. through blastoderm of an early blastula stage.

fluid-filled space, the subgerminal cavity. This causes the central zone above to look distinct and transluscent, called area pellucida. In contrast, the surrounding ring-like marginal or peripheral zone of blastoderm, lying directly on the surface of yolk, looks opaque and white and is known as the area opaca. Area pellucida later forms the body of the embryo proper, while the area opaca forms the accessory extra-embryonic structures, such as the yolk sac. Some large yolky cells of uncertain origin develop on the floor of subgerminal cavity but soon disappear (Fig. 3). According to some workers, this stage corresponds to the blastula stage of amphioxus and frog. Subgerminal cavity is regarded equivalent to blastocoel, yolky cells equivalent to megameres and central cells to micromeres. However, the subgerminal cavity is not a true blastocoel so that this stage in chick is called a pseudoblastula.

Presumptive Areas Before gastrulation, the multi-layered blastoderm of area pellucida becomes rearranged to form a single layer of cubical epithelial cells, called epiblast or ectomesoderm. Various authors (Jacobson, Pasteel, Spratt,) have mapped on this layer the prospective or presumptive areas which will later form the main embryonic structures. However, their exact locations and limits are not so well-defined as in

neural lateral

epidermal ectoderm

area

endoderm hypoblast

B

Fig. 4. Fowl. Presumptive areas In chick blastoderm before gastrulation. A-Surface view. B-In M.L.S.

·the early gastrula of Amphibia (Fig. 4). Starting at the future posterior end of embryo, the presumptive areas are as follows : (1) A small disc of endode rm. It is not shown in some fate maps because in early stages of gastrulation, the prospective endoderm cells migrate rapidly downwards into subgenninal cavity. (2) A broad rear band of lateral mesoderm. (3) A small area of prechordal mesoderm flanked on either side by somitic mesoderm. (4) A narrow band of notochordal mesoderm. (5) A large crescentic area of neural plate or neural ectoderm. (6) A still larger area of embryonic or epidermal ectoderm. All these presumptive areas lie within area pellucida. Embryonic ectoderm is continuous with extra-embryonic ectoderm on area opaca, which covers extra-embryonic endoderm.

[ 655

Development of Chick (Fowl)

Gastrulation

POSTERIOR END

ANTERIOR END

large yolky cells which will move into hypoblast by ingression

Endoderm formation. Gastrulation begins even before laying of eggs. It involves the formation of endoderm so that the monoblastic embryo or A 0::,.>y-{SJV''f("-i blastula is converted into diploblastic or two-layered gastrula. There is disagreement in different workers about the actual mode of concrescence of prospective . gastrulation, that is, format10n of endoderm in chick embryo. There is no invagination of prospective endoderm through a blastopore as 8 ,t, 1 ~. found in amphioxus and frog. From near the ' ... ~:" . .:' "~~i~~SUbg~rmin~1 space posterior end of embryo, prospective endodermal cells migrate down (involution) into the epiblast (ectoderm) subgerminal cavity, forming a coherent sheet ~~5:ID2:i~ throughout the area pellucida. There is also, to C some extent, a sinking down or ingression of individual yolk laden ectoderm cells thus adding to the thickness of blastoderm. The latter now splits hypoblast (endoderm) yolk (delamination) anteriorly into outer epiblast or ectoderm and inner hypoblast or endoderm. The Fig. 5. Fowl. Blastoderm in L.S. showmg gastrulation or successIve stages of endoderm formation. A-Large yolk cells epiblast contains the prospective neural plate, intermixed with smaller cells. B--Concrescence and notochord, mesoderm and ectoderm (Fig. 5). -

blood capillaries

connective / tissue

ciliated columnar epithelium

cllated columnar cells

blood vessel

B

A Fig. 22. A-T.S. lung of frog. 8-A portion of T. S. magnified.

moist. The inner edges of septa bear tall ciliated columnar epithelial cells.

Lung of Rabbit The mammalian lung is a pink-coloured, soft, spongy, highly elastic and highly vascular organ (Fig. 23). It lies enclosed within a special lateral air-tight compartment of thoracic cavity, the pleural cavity. The lung is invested by a fold of coelomic epithelium or visceral peritoneum. Histologically, a bronchus entering the lung repeatedly divides into finer branches, the bronchioles, each terminating into air sacs bearing numerous small hemispherical hollow projection or alveoli which present a honey-combed appearance or look like a bunch of miniature grapes. The alveoli represent the ultimate structural and physiological units of lung providing the great surface area for gaseous exchange. Extremely thin walls of air sacs and alveoli are made of a single layer of moist squamous epithelial cells and covered by a close network of capillaries, derived

12.J1/-~L-

ciliated columnar epithelial cells pulmonary capillaries

squamous epithelium

Fig. 23. T. S. portion of a mammalian lung through a bronchiole.

from branches of pulmonary artery and vein. A branchiole is lined with ciliated columnar epithelium, resting on a basement membrane, while its wall is made of elastic connective tissue and circular smooth muscle fibres. Branches of bronchus and alveoli are embedded in considerable connective tissue contammg arteries, veins, lymphatics, nerves and smooth muscle fibres.

Organ Histology of Frog & Rabbit

[ 693

perichondrium

capsule

homogeneous unless specially treated to show very fine collagen fabrils. The cells which produce matrix are called chondroblasts, with deeply stained nuclei. They lie inside fluid-filled cavities in matrix, called lacunae. They are surrounded by more deeply stained matrix called capsules. Chondroblasts are capable of mitotic cell division and usually occur in small groups of twos and fours. The mature catilage cells are known as chondrocytes.

Bone of Frog

~~"""''''';:::''''Iacunae

Fig. 24. T S. Hyaline cartilage.

Hyaline Cartilage Cartilage is a type of skeletal connective tissue. Hyaline cartilage is the commonest type (Fig. 24). It forms the whole of the adult skeleton of cartilaginous fishes (e.g. dogfish). In frog it occurs at the ends of limb bones and sternum, in hyoid apparatus and supra-scapula, etc. In mammals (e.g. rabbit) it is found in nose, ears, larynx, trachea, bronchi, xiphisternum, sternal ribs and at the ends of long bones. Peripherally, the cartilage is surrounded by a tough fibrous connective tissue sheath, called perichondrium, containing blood vessels. Often there is no sharp distinction between perichondrium and substance of cartilage. The ground substance or matrii chiefly consists of a tough, bluish-coloured, translucent glossy substance known as chondrin. Under microscope it appears

Unlike cartilage, the bone is a hard and rigid connective tissue of great strength. The intercellular substance or matrix of bone consists of an organic component, the ostein or ossein, and inorganic component including calcium and magnesium phosphates, calcium carbonate, and some sodium chloride. Ossein is in the form of collagen fibres which are somewhat difficult to detect. They yield gelatin on boiling. If a bone is burned, it becomes brittle as its organic matter is destroyed. A thin section of a typical limb bone, such as the femur of frog, shows a large central marrow cavity (Fig. 25). It is filled with a soft adipose or fatty tissue and blood vessels and termed the bone marrow. The outer bone surface is covered by a two-layered sheath called periosteum. Its outer layer consists of dense connective tissue containing blood vessels, lymphatic vessels and nerves that enter the bone. The inner layer consists of bone-forming cubical cells, called outer periosteum

~~ 4~ ~ ~ >+0 ~ >0.-~outer . "*' ~_~ __________ .""" ~

outer\'-~-~--~-~--!'-;;",\

osteoblasts

. '''J;>-

" :f5:~--:::-;t: ~;/lamellae . ~ ~ - -k . - .---- ----'"' '" osteocytes .- )'it'.""... ......... Inner \.~-, ......... "'" bone ~~::-~~~ inner layers ~.,..• 0, '. osteoblasts

bone layers

a

endosteum

A

a

oC

0

•

bone marrow

B

Fig. 25. Bone of frog. A-T. S. of decalcified femur. B-A part of decalcified bone magnified. C-A part of ground dried or calcined bone

Organ Histology of Frog & Rabbit

694] osteoblasts, arranged like an epithelium. The central marrow cavity is similarly lined by a more delicate connective tissue sheath, called endosteum, containing the inner layer of bone-forming osteoblasts. Between periosteum and endosteum, the matrix occurs in thin concentric layers or lamellae around the central marrow cavity. Between adjacent lamellae are numerous small cavities called lacunae. A lacuna gives off numerous fine branching tubules, called canaliculi. These extend in all directions, those of neighbouring lacunae opening into one another. The lacunae are occupied by bone cells called osteocytes. They differ from cartilage cells in giving off fine branching protoplasmic processes which unite with those of neighbouring cells through canaliculi, forming a continuous protoplasmic network. Thus food and oxygen can reach from peripheral blood vessels in periosteum to cells spread throughout the matrix. A ground section of dried or calcined bone seen under the microscope does not have osteocytes and their processes. Instead the lacunae and canaliculi appear black because of highly refractive air imprisoned by them. On the other hand, the section of a decalcified bone shows the osteocytes but their processes and canaliculi remain inconspicuous and often invisible.

periosteum periosteal lamellae interstitial lamellae adjacent Haversian system

The gross structure of a long bone of mammals is similar to that of frog. Periosteum is the thin, outer connective tissue sheath surrounding the bone (Fig. 26). The central marrow cavity contains yellow marrow in the shaft (diaphysis) and red marrow at the ends (epiphyses). Yellow marrow consists primarily of fat and produces white corpuscles. Red marrow consists primarily of cells called myelocytes and produces red corpuscles. However, the arrangement of lamellae varies from that of frog. Matrix between periosteum and endosteum is perforated by a series of fine channels called Haversian canals after the English anatomist, Clopton Havers. These run more or less parallel to the long axis of the bone. Each Haversian canal contains an artery, a vein, a lymphatic vessel, nerve fibres, occasional fat cells, flattened osteoblasts and a small amount of areolar connective tissue. Each canal is surrounded by concentric layers of bone or lamellae. Minute spaces or lacunae housing bone cells or osteocytes also form concentric rings alternating with the lamellae. Fine protoplasmic processes of osteocytes extend through canaliculi and freely anastomose linking cells to one another and to Haversian canal. Concentric lamellae enclosing an Haversian canal constitute a Haversian system, which

H=~8

(.~

blood

~~~o

~~. osteocytes

A

Bone of Rabbit

bone lamellae

·B

C

Fig. 26. Bone of rabbit. A-T S. of a part of dried or calcined bone B-Decalcified. C-A haversian system magmfied

[695

Organ Histology of Frog & Rabbit

presents a very characteristic form in the section of a mammalian bone. Haversian canals are interconnected by transverse Volkmann canals which contain blood vessels but are not surrounded by concentric lamellae. The interstices or intervals between adjoining Haversian systems are occupied by irregular interstitial lamellae and cells.

BLOOD CELLS FROG'S BLOOD

§@),', . .:."':, ERYTHROCYTESO~ ',,',.;tift,

Q i

,

HUMAN BLOOD

",

I

(red cells)

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"

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~' y~

nucleus

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II LEUCOCYTES (white cells)

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~~ ~ ,,~

A-GRANULOCYTES (polymorphonuclear leucocytes)

Blood Blood is a liquid tissue in which round cells (corpuscles), entirely separate from one another, float in a pale straw-coloured intercellular fluid, the plasma. Blood cells or corpuscles are of two types : red and white (Fig. 27). Besides, there are cell fragments or blood plates or platelets. 1. Erythrocytes (red corpuscles). These are so called because they contain the red pigment haemoglobin. In lower vertebrates (e.g. frog), red corpuscles are oval, thin, biconvex discs, each having a large, oval, central nucleus. In mammals (e.g. rabbit), red corpuscles are circular, thin, biconcave discs without nuclei. Red corpuscles of rabbit are 811 in diameter, while those of frog are much larger, about 2511 in length. They are formed in the red bone marrow and serve as oxygen carrier. 2. Leucocytes (white corpuscles). These are so called because they are colourless. They differ from erythrocytes in lacking permanent shape, in retaining nuclei, but having no haemoglobin. They are formed in bone marrow, spleen and lymphoid tissue and are phagocytic in nature. There are 5 or 6 different kinds of leucocytes. They belong to two general types, granulocytes and agranulocytes. (a) Granulocytes (polymorphonuclear leucocytes). The nucleus is strongly lobulated and variable in shape. The cytoplasm contains prominent granules with specific staining property. Accordingly, there are 3 types of granulocytes. Granules of acidophils or eosinophils stain pink with acid dyes, basophils stain blue with basic ,dyes, whereas neutrophils take up both the acid and basic dyes. (b) Agranulocytes (mononuclear leucocytes). They lack the cytoplasmic granules. Lymphocytes

~ "

" .:'

, ", : ,",',

I NEUTROPHILS I multilobed nucleus

IEOSINOPHILS I bilobed

IBASOPHILS I nucleus--

G~. •• ~"'",

'

.. ~~~:".

B-AGRANULOCYTES (mononuclear leucocytes)

ILYMPHOCYTES I nucleu

I MONOCYTES I III THROMBOCYTES (blood platelets)

Fig. 27. Blood cells of frog and rabbit.

have a large spherical nucleus which nearly fills the cell. Monocytes are relatively larger cells with an oval to kidney-shaped nucleus. 3. Platelets (thrombocytes). They are not considered proper cells but only cell fragments. They are very small bodies without nucleus in mammals. In other vertebrates (e.g. frog), they are spindle-like and nucleated. Blood platelets function in connection with clot-formation.

Organ Histology of Frog & Rabbit

696 1

Spleen Spleen is the largest lymphoid gland in the body of vertebrates (Fig. 28). It is surrounded by a definite fibromuscular capsule covered externally by a layer of flattened cells of serosa or visceral peritoneum. From the capsule extend, usually at right angles, numerous fibro-muscular strands or trabeculae into the soft spongy parenchyma or splenic pulp which is supported by reticular tissue and divided into lobules. The splenic pulp is composed of red and white pulps. The red pulp, forming the bulk, contains mostly free erythrocytes which give it red colour. Red pulp also comprises irregularly shaped pulp cords composed of erythrocytes, leucocytes, reticular cells, macrophages and splenocytes. Fairly large and irregular spaces also occur, called venous sinuses or terminal veins. The red pulp also contains scattered patches of splenic nodules forming white pulp. They look white in fresh tissue but darker than red pulp in a stained preparation. They are formed by accumulations of lymphocytes, large phagocytic macrophages or histiocytes and reticular cells, and usually posssess an eccentric artery or arteriole. Spleen also contains trabecular arteries, veins and nerves. Spleen serves as a reservoir of erythrocytes. Its macrophages destroy worn out old erythrocytes, while lymphocytes are concerned with the production of antibodies.

visceral

npritnn,,,,, '~~~fibrous

capsule

venous

Fig. 28. T. S. of a small portion of spleen.

dorsal septum

subarachnoid space

ventral septum

central canal

Fig. 29. T. S. Spinal cord offrog.

Spinal Cord The histological structure of spinal cord, as seen in a transverse section, is the same in all the vertebrates (Figs. 29 & 30). Like brain, it is surrounded by two protective and fibrous membranes (meninges). The thick outer meninx is called duramater, while the thin, inner and vascular one, the piamater. Between the two lies the subarachnoid fluid which protects the spinal cord from external shocks. Two deep clefts or grooves, the dorsal and ventral median fissures, divide the spinal cord into right and left lateral halves. The lumen at the centre of the cord is called the central canal. It is lined with ciliated columnar epithelium (ependyma) and contains the

Fig. 30. T. S. Spinal cord of rabbit.

[ 697

Organ Histology of Frog & Rabbit cerebrospinal fluid. Two types of nerve tissue make up the spinal cord, inner gray matter and outer white matter, so called because of their appearance in fresh condition. The central rectangular or H-shaped gray matter consists of nerve cells (neurons), non-medullated fibres, neuroglia . and blood vessels. The central canal divides the gray commissure into dorsal and ventral portions. The gray matter is projected at the corners into dorsal and ventral homs to which roots of spinal nerves are attached. The outer or peripheral white matter mainly consists of medullated nerve fibres. Their bundles are arranged on either side into dorsal, lateral and ventral funiculi. The spinal cord conducts impulses to and from brain and also controls the reflex activity.

Kidney of Frog Kidney of adult frog is an opisthonephros or mesonephros (Fig. 31). It is invested by a fibrous connective tissue capsule, which is covered only

ventrally by visceral peritoneum of flat cells. In a T.S., kidney shows a large number of uriniferous tubules cut in various planes and formed by cuboidal epithelium. Malpighian bodies lie close to the ventral border while collecting tubules near the dorsal border of kidney. A Malpighian body includes a bunch of blood capillaries, called glomerulus, enclosed by a close double-walled cup, the Bowman s capsule. In an ordinary preparation, the walls of capillaries forming the glomerulus are not distinct, but the blood cells are easily seen. Walls of Bowman's capsule consist of squamous epithelial cells. Ureter may be seen close to the lateral border of kidney, cut transversely and lined by columnar epithelium. Tubules and capsules are embedded in connective tissue which also contains muscle and nerve fibres and renal arteries and veins cut at several places. Ventral surface of kidney shows several ciliated funnels, called nephrostomes, which communicate coelom with the veins of kidney.

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Bowman's, capsule